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UNIVERSITY CM- 
ILLINOIS LIBRARY 
AT URBANA-CHAMPAIGN 
NATURAL HIST 8UBV 



FIELDIANA 
Botany 

Published by Field Museum of Natural History 

. 



FLORA OF PERU 

I. FRANCIS MACBRIDE 
AND COLLABORATORS 

FAMILY COMPOSITAE: PART III 

GENUS MIKANIA - - TRIBE EUPATORIEAE 
WALTER C. HOLMES 
SIDNEY McDANIEL 



July 29, 1982 
Publication 1333 



FLORA OF PERU 

FAMILY COMPOSITAE: PART III 

GENUS MIKANIA TRIBE EUPATORIEAE 



FIELDIANA 
Botany 

Published by Field Museum of Natural History 



New Series, No. 9 



FLORA OF PERU 

J. FRANCIS MACBRIDE 
AND COLLABORATORS 



FAMILY COMPOSITAE: PART III 

GENUS M1KAN1A TRIBE EUPATORIEAE 

WALTER C. HOLMES 
Institute for Botanical Exploration 
Mississippi State, Mississippi 
Department of Biological Science 
Northwestern State University of Louisiana 
Natchitoches, Louisiana 

SIDNEY McDANIEL 
Institute for Botanical Exploration 
Mississippi State, Mississippi 
Department of Biological Sciences 
Mississippi State University 
Mississippi State, Mississippi 



Accepted for publication November 10, 1981 
July 29, 1982 
Publication 1333 



Library of Congress Catalog Card No.: 80-66384 

ISSN 0015-0746 
PRINTED IN THE UNITED STATES OF AMERICA 



CONTENTS 

List of Illustrations vii 

Mikania 1 

Keys to Species 3 

1. Mikania angularis 8 

2. Mikania aquaria 8 

3. Mikania areolata 8 

4. Mikania arthroclada 9 

5. Mikania aschersonii 9 

6. Mikania banisteriae 11 

7. Mikania brachyphylla 13 

8. Mikania brevifaucia 13 

9. Mikania brooksii 13 

10. Mikania bulbisetifera 14 

11. Mikania bullata 14 

12. Mikania carnosa 15 

13. Mikania chimborazensis 15 

14. Mikania congesta 16 

15. Mikania conglomerata 16 

16. Mikania cordifolia 17 

17. Mikania crassifolia 18 

18. Mikania cutervensis 18 

19. Mikania cuzcoensis 19 

20. Mikania decora 19 

21. Mikania desmocephala 21 

22. Mikania dudleyi 21 

23. Mikania ecuadorensis 22 

24. Mikania featherstonei 22 

25. Mikania filicifolia 22 

26. Mikania glandulifera 23 

27. Mikania guaco 23 

28. Mikania haenkeana 24 

29. Mikania hexagonocaulis 26 

30. Mikania hookeriana 26 

31. Mikania huanucoensis 27 

32. Mikania incasina 28 

33. Mikania infesta 28 

34. Mikania inornata 28 

35. Mikania iquitosensis 29 

36. Mikania jelskii 29 

37. Mikania juninensis 30 

38. Mikania klugii 30 

39. Mikania lanceolata 30 

40. Mikania lancifolia 31 



VI 



41. Mikania lasiopoda 31 

42. Mikania leiostachya 32 

43. Mikania lindleyana 32 

44. Mikania lutescens 32 

45. Mikania marinii .'. 33 

46. Mikania mathewsii 33 

47. Mikania mazanensis 34 

48. Mikania megalophylla 35 

49. Mikania micrantha 36 

50. Mikania microptera 39 

51. Mikania montana 39 

52. Mikania ochroleuca 40 

53. Mikania oreopola 40 

54. Mikania parvicapitulata 41 

55. Mikania parviflora 41 

56. Mikania pendula 42 

57. Mikania psilostachya 42 

58. Mikania pycnadenia 44 

59. Mikania rimachii 44 

60. Mikania rugosa 46 

61. Mikania shushunensis 47 

62. Mikania simpsonii 48 

63. Mikania speciosa 48 

64. Mikania stereodes 49 

65. Mikania stuebelii 49 

66. Mikania stygia 50 

67. Mikania szyszylowiczii 50 

68. Mikania tambillensis 50 

69. Mikania ternata 51 

70. Mikania trachodes 52 

71. Mikania turbaricola 53 

72. Mikania vargasii 53 

73. Mikania violaefolia 53 

74. Mikania vitifolia 54 

75. Mikania weberbaueri 56 

76. Mikania woytkoivskii 56 



LIST OF ILLUSTRATIONS 

1 . Mikania aschersonii Hieron 10 

2. Mikania decora Poepp 20 

3. Mikania guaco H. & A 25 

4. Mikania mathewsii B. L. Robinson 34 

5. Mikania megalophylla Holmes & McDaniel 35 

6. Mikania micrantha HBK 37 

7. Mikania psilostachya DC 43 

8. Mikania rimachii Holmes & McDaniel 45 

9. Mikania shushunensis Holmes & McDaniel 47 

10. Mikania ternata (Veil.) B. L. Robinson 51 

11. Mikania vitifolia DC 55 



vii 



MIKANIA 

Mikania Willd., Sp. PI. 3: 1742. 1803. TYPE: M. scandens (L.) Willd. (Eupatorium 
scandens L.). Name and type conserved. 

Carelia A. L. Jussieu ex Cav., An. Ci. Nat. 6: 317. 1802 (1803), not Carelia G. Pontedera 
ex Fabricus 1759. TYPE: C. satureiaefolia Cav. = Mikania amara Willd. 

Corynanthelium Kunze, Linnaea 20: 19. 1847. TYPE: C. moronoa Kunze. 

Moronoa hort. ex Kunze, Linnaea 20: 19. 1847, nom. nud. pro syn. 

Morrenia hort. ex Kunze, Linnaea 20: 19. 1847, nom. nud. pro syn. TYPE: M. odorata 
Kunze, nom. nud. 

Willoughbya Neck, ex Kunze, Rev. Gen. PI. 371. 1891. TYPE: Eupatorium scandens L. = 
Mikania scandens (L.) Willd. 

Willughbaeya Neck., Elem. 1: 82. 1790, nom. nud. 

Herbs or woody perennials, twining or somewhat shrubby, rarely erect; stems round or 
hexagonal. Leaves opposite, rarely verticillate, usually petiolate, simple, entire, toothed 
or rarely divided, pinnately or palmately veined. Inflorescence paniculate, the ultimate 
segments spicate, racemose, corymbose, cymose, glomerate, or thyrsiform. Heads 
homogamous, 4-flowered, rarely 3- or 5-flowered; involucral bracts 4, equal, mostly ob- 
long, with or without a fifth outermost one (exterior bract or bracteole); receptacle small, 
naked, glabrous; corollas white, pink-purple, green, pale greenish yellow (to reportedly 
pale yellow in M. lutescens), tubular, the tube slender, sometimes slightly enlarged at the 
base by the enclosed nectary, the throat gradually or suddenly enlarged, turbinate, cylin- 
dric, or campanulate, corolla teeth 5, equal, deltoid or lance-linear, usually shorter than 
but sometimes exceeding the length of the throat; anthers basally rounded or more rarely 
emarginate or obscurely subsagittate, the terminal appendage ovate or oblong; style 
branches elongate, clavellate, or filiform. Achenes pentagonal, prismatic or tapering 
downward; pappus bristles capillary, numerous in 1 or rarely 2 series, slightly scabrid or 
barbellate, rarely smooth, free or very slightly connate into a narrow ring at the base, 
white, pale buff, or fulvous. 

Mikania (Compositae-Eupatorieae) is well represented in Peru, this present 
work recognizing 76 species. This number is only exceeded in Brazil, with five 
times the area of Peru, where Barroso (1959) reports 152 species. The Peruvian 
species have received no complete taxonomic treatment since the work of 
Robinson (1922) on the Mikania species of northern and western South America. 
Robinson recognized 37 species, many based upon description alone and 
numerous others being superfluous synonyms based upon the premise that one 
species was unlikely to occur in two widely separated areas, i.e., a phytogeo- 
graphic species concept. Since then most work has consisted of descriptions of 



This study was assisted, in part, by National Science Foundation Grant DEB- 79-05078 
(Alwyn H. Gentry, principal investigator). 



FIELDIANA: BOTANY 

proposed species, making determinations difficult, if not impossible, without 
comparison with types. This present paper is intended to provide keys, de- 
scriptions, specimen citations, and nomenclatural clarification of all known 
Peruvian species of Mikania. 

This work is the result of herbarium and field studies begun in 1973 and is 
part of a continuing effort to revise the entire genus. Material was available 
from 20 institutions and we are greatly indebted to the curators of those in- 
stitutions for the loan of specimens which included types of most of the Peru- 
vian species of Mikania. Two herbaria not included in Index Herbariorum (ed. 
6) are Herbarium Amazonense (AMAZ), Universidad de la Amazonia 
Peruviana, Iquitos, Peru, and Institute for Botanical Exploration (IBE), Mis- 
sissippi State, Mississippi. 

Mikania is a well-marked genus of mostly perennial twiners with opposite to 
rarely verticillate leaves. Heads are singularly uniform, each possessing four 
flowers enclosed by four involucral bracts that are usually subtended by an 
exterior bract. This uniformity within the heads is not reflected in the disposi- 
tion of the heads into an inflorescence of a secondary nature (capitulescence) 
which may be spicate, racemose, thyrsoid, corymbose (including temately 
branching corymbs), or glomerate. In this work inflorescence actually r fUB3riJ n to 
capitulescence, which has been standard usage in the genus. The nature' of the 
capitulescence has been the major basis of segregation of the genus into sec- 
tions. These sections are not followed, due to intergradation, and sucrrdivision 
is best postponed pending further studies and a treatment of the entiajgenus. 

Although the genus is probably one of the most easily recognized^and most 
uniform of the Compositae-Eupatorieae, difficulty is often encountered in 
specific delimitation. Aside from the large number of species and sparsity of 
current works, this difficulty is due, in large part, to several highly polymorphic 
species "complexes" that have been discussed by Robinson (1934) and Steyer- 
mark (1953). When portions of these "complexes" are examined for a limited 
area, they may well appear to be good species. Critical study over a wider area, 
however, shows these species to be connected by intermediates. Normally the 
major differences encountered concern the type or amount of pubescence and 
leaf characters (shape, texture, etc.), which, in one example, have been shown 
to vary on the same plant and are highly unreliable for specific delimitation (see 
Steyermark, 1953). Among the polymorphic species are Mikania banisteriae DC., 
M. hookeriana DC., M. vitifolia DC., M. micrantha H.B.K. and M. parviflora 
(Aubl.) Karst. Certainly more names will be added to this list in future studies. 
In these groups, as well as certain others, the species concept followed is similar 
to that of the treatment by Steyermark (1953) of M. banisteriae. For further 
information on these complexes, refer to the taxonomic treatment. 

In construction of the keys, an attempt was made to utilize the most obvious 
character, mostly macroscopic, that would separate the species, usually leaf 
characters, type of secondary inflorescence, or head size. Between closely re- 
lated species, use of a 10 x lens was often necessary, particularly for characters 
of involucral bracts and corollas. In this work, the corolla tube is the un- 
expanded portion and the throat the expanded portion less the teeth. The 
peduncles supporting the individual heads are referred to as "pedicels," which 
is common usage in this genus. Leaf measurements are for the blade only, 
petiole length is given separately. Heads are measured from the base of the 
involucre to the tips of the corolla tube. A conscious effort was made to include 



MACBRIDE: FLORA OF PERU 3 

key characters with little or no leeway in interpretation; however, this was not 
always possible. In instances where a character could be interpreted in more 
than one way, the species involved should key under both leads. 

REFERENCES 

BARROSO, G. M. 1959. Mikaniae do Brasil. Arquiv. Jard. Bot. Rio de Janeiro 16: 239-333. 
CABRERA, A. L. 1953. Compuestas Peruanas nuevas o criticas. Bol. Soc. Argent. Bot. 5: 

37-50. 
HOLMES, W. C, AND S. MCDANIEL. 1979. Notes on Mikania (Compositae) V. Phytologia 

41: 183-197. 

ROBINSON, B. L. 1920. Further diagnoses and notes on tropical American Eupatorieae. 
Contr. Gray Herb. 61: 1-30. 

. 1922. The Mikanias of northern and western South America. Contr. Gray Herb. 
64: 21-116. 

-. 1924. Records preliminary to a general treatment of the Eupatorieae IV. Contr. 



Gray Herb. 73: 1-31. 

-. 1926. Records preliminary to a general treatment of the Eupatorieae VI. Contr. 



Gray Herb. 77: 1-62. 

-. 1928. Records preliminary to a general treatment of the Eupatorieae VII. Contr. 



Gray Herb. 80: 1-42. 

-. 1934. The variability of two wide-ranging species of Mikania. Contr. Gray Herb. 



104; 49-55. 

STEYERMARK, J. A. 1953. Botanical exploration in Venezuela III. Fieldiana, Bot. 28: 648- 
658. 

KEY TO GROUPS 

1. Leaves parted or divided to midvein I. 

1. Leaves entire, serrate, crenate, etc., but not parted to midvein. 

2. Heads in subspherical somewhat dense glomerules II. 

2. Heads in spikes, racemes, panicles, or corymbs, but not in dense subspherical 
glomerules. 

3. Heads in spikes or racemes ni. 

3. Heads in panicles or corymbs. 

4. Heads sessile or nearly so and regularly in groups of 3's at the tips of 

branches IV. 

4. Heads pedicellate, not in groups of 3's at tips of branches. 

5. Leaves palmately veined V. 

5. Leaves pinnately or subpinnately veined VI. 



KEYS TO SPECIES 

GROUP I 

1. Leaves 2-3-pinnate, leaflets less than 1 cm in diameter 25. M. filicifolia. 

1. Leaves palmately divided; leaflets usually wider than 1 cm 69. M. ternata. 

GROUP II 

1. Leaves cordate at base. 

2. Heads in a congested spiciform continuous or more or less interrupted glom- 

erule 28. M. haenkeana. 

2. Heads in a very dense subspherical glomerule. 

3. Exterior bract ca. Vz the length of the involucre, leaves semicoriaceous, leaf bases 

truncate to at most subcordate 15. M. conglomerata. 

3. Exterior bract as long as or greater than the length of the involucre, leaves 

membranous, leaf bases distinctly cordate 14. M. congesta. 

1. Leaves truncate, cuneate, or rounded at base, not at all cordate. 



4 FIELDIANA: BOTANY 

4. Heads 5-6 mm long, mature cauline leaves 15 cm or more long (mature cauline 
leaves absent on many specimens). 
5. Leaves distinctly feather veined, inflorescence spicate 48. M. megalophylla. 

5. Leaves subpinnately veined from near the base, inflorescence dense, sub- 
spherical 15. M. conglomerata. 

4. Heads 8-10 mm long, mature cauline leaves usually less than 15 cm long. 

6. Leaves ovate to broadly ovate, involucral bracts distinctly and prominently 
swollen at the base, glomerule somewhat elongate 30. M. hookeriana. 

6. Leaves lanceolate to narrowly ovate, involucral bracts scarcely swollen at the 
base, glomerule spherical. 

7. Corolla ca. 4 mm long, exterior bract ovate, leaf bases truncate 

21. M. desmocephala. 

7. Corolla ca. 6-6.5 mm long, exterior bract linear (not seen in M. weberbaueri), 
leaf bases acute to obtuse. 

8. Leaf apices acute to abruptly acuminate, pappus bristles not thickened at 
tips 68. M. tambillensis. 

8. Leaf apices long attenuate, pappus bristles thickened at tips 

75. M. weberbaueri. 

GROUP III 

1. Leaves verticillate 62. M. simpsonii. 

I. Leaves opposite. 

2. Leaves clearly palmately veined (at least on mature cauline leaves). 
3. Leaf bases distinctly cordate to subcordate. 

4. Heads on pedicels ca. 5 mm long 19. M. cuzcoensis. 

4. Heads sessile or nearly so. 

5. Leaf margins conspicuously serrate, teeth ca. 1 cm apart 

56. M. pendula. 

5. Leaf margins entire. 

6. Heads ca. 5 mm long, in rather dense, somewhat elongated glomerules 

28. M. haenkeana. 

6. Heads 7-12 mm long, in rather loose elongated spikes. 

7. Heads 10-12 mm long 72. M. vargasii. 

7. Heads 7-9 mm long 2. M. aquaria. 

3. Leaf bases truncate or cuneate, but not clearly cordate or subcordate. 

8. Leaves deltoid, hastately lobed at the base, base truncate at insertion of 
petiole 52. M. ochroleuca. 

8. Leaves ovate or elliptic, not hastately lobed at base. 

9. Leaves elliptic, less than 4 cm wide 76. M. woytkowskii. 

9. Leaves ovate, 5 cm or more wide. 

10. Heads in rather dense, subspherical glomerules, base of involucral 

bracts swollen 30. M. hookeriana. 

10. Heads in elongated spikes, base of involucral bracts not swollen 

2. M. aquaria. 

2. Leaves clearly pinnately veined. 
11. Upper leaf surfaces scabrous. 

12. Heads 8-10 mm long 57. M. psilostachya. 

12. Heads ca. 5 mm long 33. M. infesta. 

11. Upper leaf surfaces not at all scabrous. 

13. Heads distinctly pedicellate. 

14. Leaves at most 5 cm long, coriaceous, involucral bracts 2.5 mm long . . 
54. M. parvicapitulata. 

14. Leaves 6-15 cm long, thin, involucral bracts 3 mm long 

20. M. decora. 

13. Heads sessile or nearly so. 

15. Leaf bases cuneate. 

16. Leaf apices rounded to blunt 67. M. szyszylowiczii. 

16. Leaf apices acute to acuminate 31. M. huanucoensis. 

15. Leaf bases obtuse to truncate. 

17. Heads in 3's and 5's at the tips of the branchlets . . .65. M. stuebelii. 
17. Heads not in 3's or 5's at the tips of the branchlets. 



MACBRIDE: FLORA OF PERU 5 

18. Mature cauline leaves greater than 25 cm long, heads antrorsely 

oriented, not perpendicular to branches of inflorescence 

48. M. megalophylla. 

18. Mature cauline leaves less than 20 cm long, heads perpendicular 

to branchlets or inflorescence. 

19. Inflorescence lax, heads remotely spaced on branchlets. 
20. Corolla tube 1-1.2 mm long 53. M. oreopola. 

20. Corolla tube ca. 2 mm long 38.. M. klugii. 

19. Inflorescence dense, heads on branchlets usually less than 

5 cm long. 

21. Corolla tube ca. 0.5 mm long 36. M. jelskii. 

21. Corolla tube 1.0 mm long or more. 

22. Leaves elliptic-ovate, plant with dark glandular 
punctations on leaves and branchlets, leaf margins 
not revolute 26. M. glandulifera. 

22. Leaves ovate, lacking glandular punctations, leaf 
margins revolute 42. M. leiostachya. 

GROUP IV 

1. Corolla teeth greater than or equal to the length of the throat 63. M. speciosa. 

1. Corolla teeth less than the length of the throat. 

2. Leaves lanceolate, ca. 4x or more longer than wide. 

3. Plant blackening upon drying, involucral bracts oblong, ca. 8 mm long 

66. M. stygia. 

3. Plant not blackening upon drying, involucral bracts spatulate, ca. 3 mm long. . . 

39. M. lanceolata. 

2. Leaves ovate, oblong, or elliptic, less than 3x longer than wide. 

4. Leaf bases conspicuously decurrent upon petiole 27. M. guaco. 

4. Leaf bases truncate, obtuse or acute to shortly cuneate, but not at all con- 
spicuously decurrent upon the petiole. 

5. Inflorescence a panicle 65. M. stuebelii. 

5. Inflorescence a corymb or heads in a somewhat dense subspherical 
glomerule. 

6. Inflorescence a very dense corymb, appearing as a rather dense sub- 
spherical glomerule 68. M. tambillensis. 

6. Inflorescence a more or less lax corymb. 
7. Petioles at most 5-6 mm long. 

8. Leaves densely punctate with reddish-orange glands, involucral 
bracts ca. 6.5 mm long 58. M. pycnadenia. 

8. Leaves at most puberulent, not punctate with reddish-orange 
glands, involucral bracts ca. 4.5 mm long 9. M. brooksii. 

7. Petioles 10 mm or more long. 

9. Stem somewhat hexagonal, exterior bract ovate, petiolate, ca. V4 or 
less the length of the involucre 29. M. hexagonocaulis. 

9. Stems terete, exterior bract obovate, somewhat petiolate, ca. Vz or 
more the length of the involucre 55. M. parviflora. 

GROUP V 

1. Inflorescence paniculate. 

2. Leaf bases cordate to hastate. 

3. Heads 4-5 mm long, in rather dense somewhat elongate spikes 

28. M. haenkeana. 

3. Heads 7-10 mm long, inflorescence lax, not at all spicate. 

4. Corolla throat campanulate, teeth about as long as throat ... .74. M. vitifolia. 

4. Corolla throat salverform-funnelform, teeth ca. 3x as long as throat 

1. M. angularis. 

2. Leaf bases cuneate to subtruncate. 

5. Corolla teeth shorter than throat. 



6 FIELDIANA: BOTANY 

6. Heads sessile, in rather dense somewhat elongated spikes, involucral bracts 
swollen at base 30. M. hookeriana. 

6. Heads with pedicels 2-6 mm long, in open panicles, involucral bracts not 
swollen at base 71. M. turbaricola. 

5. Corolla teeth longer than throat. 

7. Exterior bract about as long as involucre, branches of inflorescence gla- 
brate 34. M. inornata. 

7. Exterior bract ca. Vz as long as involucre, branches of inflorescence crisped- 

puberulent 32. M. incasina. 

1. Inflorescence corymbose. 

8. Leaf bases cuneate, truncate, obtuse, but not distinctly cordate or subcordate. 
9. Leaves broadly ovate, greater than 5 cm long. 

10. Heads 12-14 mm long 17. M. crassifolia. 

10. Heads 8-9 mm long. 

11. Leaves entire 43. M. lindleyana. 

11. Leaves denticulate 45. M. marinii. 

9. Leaves ovate-lanceolate, less than 5 cm long. 

12. Leaf bases rounded, exterior bract as long as the involucre, leaves prom- 
inently reticulate-areolate 51. M. montana. 

12. Leaf bases acute, exterior bract 2 as long as the involucre, leaves not prom- 
inently reticulate-areolate 12. M. carnosa. 

8. Leaf bases definitely cordate to subcordate. 
13. Stems distinctly hexagonal. 

14. Angles of stem riblike, not winged, plants tomentose to tomentulose 

16. M. cordifolia. 

14. Angles of stem with narrow subherbaceous wings, plants glabrate to pu- 
berulent 50. M. microptera. 

13. Stems terete, quadrangular, multistriate, costate, but not distinctly hexagonal. 

15. Leaves less than 1 cm long 7. M. brachyphylla. 

15. Leaves greater than 1 cm long, usually 3 cm or more. 

16. Corolla teeth greater than or about equaling the length of the throat. 
17. Inflorescence lax, pedicels 5-15 mm long, exterior bract borne well 

beneath head 47. M. mazanensis. 

17. Inflorescence not lax, pedicels less than 5 mm long, exterior bract 
borne at summit of pedicel. 
18. Leaves triangular, corolla teeth ca. 3x or more as long as 

throat 8. M. brevifaucia. 

18. Leaves broadly ovate, corolla teeth about as long as the 

throat 59. M. rimachii. 

16. Corolla teeth less than the length of the throat. 
19. Heads 4-7 mm long. 

20. Plant velutinous, pappus bristles 20-22, smooth 

23. M. ecuadorensis. 

20. Plant glabrate to puberulent, not velutinous, pappus bristles 
30-35, barbellate 49. M. micrantha. 

19. Heads 10-16 mm long. 

21. Leaves 5 cm or less long, triangular, margins dentate, heads 
10-11 mm long. 

22. Leaves membranous, not glandular, stems not purplish 
44. M. lutescens. 

22. Leaves subcoriaceous to somewhat fleshy, glandular, stems 
purplish 73. M. violaefolia. 

21. Leaves greater than 5 cm long, margins entire to serrate or den- 
tate, heads 12-16 mm long. 

23. Corolla teeth less than 1 mm long 3. M. areolata. 

23. Corolla teeth 1.5 mm or more long. 

24. Leaves coarsely dentate, corolla tube ca. 1.5 mm long, 
throat ca. 6.5 mm long 18. M. cutervensis. 



MACBRIDE: FLORA OF PERU 7 

24. Leaves serrate-crenate, corolla tube and throat about 
equal in length (ca. 5 mm) 13. M. chimborazensis. 

GROUP VI 

1. Inflorescence corymbose. 

2. Heads 12-13 mm long, leaves ovate with cordate bases, stems densely puberu- 

lent 24. M. featherstonei. 

2. Heads 8-10 mm long, leaves lanceolate with cuneate to obtuse bases, stems gla- 
brous to villous when young. 

3. Inflorescence a rather dense corymb, leaf apices long attenuate 

75. M. weberbaueri. 

3. Inflorescence an open corymb, leaf apices acuminate 5. M. aschersonii. 

1. Inflorescence paniculate. 

4. Corolla throat ca. 2x as long as teeth or tube and throat indistinguishable. 

5. Leaves at most ca. 1.5 cm long, margins crenate-serrate, plants densely reddish 

pubescent 4. M. arthroclada. 

5. Leaves usually 3 cm or more long, margins not crenate-serrate, plant not reddish 
pubescent. 
6. Involucral bracts 5 mm or more long. 

7. Leaves coriaceous, margins revolute, often coarsely and irregularly ser- 
rate, achenes densely glandular 71. M. turbaricola. 

7. Leaves membranous, margins entire, achenes nonglandular. 

8. Leaf bases rounded to truncate, corolla throat campanulate, teeth del- 
toid 35. M. iquitosensis. 

8. Leaf bases cuneate, corolla throat and tube indistinguishable, teeth 

lanceolate 61. M. shushunensis. 

6. Involucral bracts 4.5 mm or less long (usually less than 4 mm long). 

9. Corolla tube 1 mm or less long, ca. V* the length of the throat, leaves 

scabrid above 64. M. stereodes. 

9. Corolla tube longer than 1 mm, ca. % the length of the throat, leaves 
smooth above. 

10. Leaf bases acuminate 46. M. mathewsii. 

10. Leaf bases cordate (bracteal leaves subcordate to truncate) 

22. M. dudleyi. 

4. Corolla throat ca. the same length as the teeth. 
11. Involucral bracts 4-4.5 mm long. 

12. Leaves lanceolate, margins prominently callose-denticulate 

40. M. lancifolia. 

12. Leaves ovate, elliptic, or lanceolate, margins entire or at most remotely and 
obscurely callous-denticulate. 
13. Involucral bracts glabrate, apices rounded, exterior bract ovate, borne 

slightly beneath the involucre 6. M. banisteriae. 

13. Involucral bracts pilose-hispid, tawny pilose, tomentose, or villous; 
apices acute to rounded; exterior bract linear to lanceolate to oblong, 
bome at the summit of pedicel. 

14. Stems somewhat hexagonal 60. M. rugosa. 

14. Stems terete. 

15. Stems scabrous 70. M. trachodes. 

15. Stems lanate to tomentose. 

16. Heads 9-10 mm long, exterior bract narrowly ovate 

10. M. bulbisetifera. 

16. Heads 7-8 mm long, exterior bract linear to oblanceolate . . 

11. M. bullata. 

11. Involucral bracts 3-3.5 mm long. 

17. Corolla tube abruptly expanded into a broadly campanulate throat, cauline 

leaves with cordate bases 41. M. lasiopoda. 

17. Corolla tube gradually expanded into a turbinate throat, cauline leaves with 
rounded to obtuse bases 37. M. juninensis. 



8 FIELDIANA: BOTANY 

1. Mikania angularis H. & B., PL Aeq. 2: 87. 1809. TYPE: Ecuador, Loja, near 
Gonzanama, Humboldt & Bonpland 3434 (P, holotype, not seen, F, photo). 

M. laxa DC, Prodr. 5: 200. 1836. TYPE: Peru, Cuchero, Poeppig 1218 (G-DC, holotype, 

not seen, IDC microfiche 800. 822: I. 5; F, photo). 
M. armigera Poepp. in Poepp. & Endl., Nov. Gen. Spec. PI. 3: 52. 1845. TYPE: Peru, 

Pampayaco, Poeppig D (W, holotype, not seen, GH, photo; F, isotype). 
Not including M. salviaefolia Gardn. considered synonymous with M. armigera by 

Baker in Mart., Fl. Bras. 6(2): 250. 1876. 

Somewhat woody twiners; stems terete, glabrate to somewhat rusty-puberulent. 
Leaves triangular-hastate, to ca. 13 cm long and 13 cm wide, margins entire, apices acute 
to acuminate, bases subtruncate to cordate, with 2 divergent acute to acuminate basal 
angles at the sides, surfaces glabrate to somewhat puberulent, 3-5-nerved from the base; 
petioles 5-8 cm long, slender, puberulent. Inflorescence a panicle, to ca. 30 cm long; 
branchlets weakly angular, rusty puberulent; pedicels 2-5 mm long, rusty puberulent. 
Heads 9-10 mm long; exterior bracts ovate-lanceolate, ca. Va the length of the involucre; 
involucral bracts oblong, ca. 6mm long, glabrate, apices rounded; corollas 5-5. 5 mm long, 
white, tube ca. 3.5 mm long, thin, throat abruptly expanded, ca. 0.3 mm long, teeth 
ovate-oblong, 1.5-1.7 mm long. Achenes ca. 2.7 mm long, grayish brown, angles up- 
wardly scabrid; pappus bristles 30-35, white, ca. 5 mm long, barbellate, thickened at the 
tips. 

Mikania angularis, of Peru, Ecuador, and Colombia, is very similar to M. 
viti folia, but with corolla teeth 3x or more the length of the throat. Mikania 
vitifolia has corolla teeth about the same length as the throat. Comparison of M. 
laxa with M. angularis showed the two to be conspecific (see Holmes & 
McDaniel, 1979). 

CUZCO. Paucartambo: Piuipiui, 720 m, Vargas 17793 (US). HUANUCO. 
Huamalies: Monzon River, Weberbauer 3640 (GH). JUNIN. La Merced, 
Hacienda Schunke, Macbride 5290 (F, GH, US). 

2. Mikania aquaria B. L. Robinson, Contr. Gray Herb. 73: 21. 1924. TYPE: Peru, 
Junin, La Merced, Macbride 5242 (F, holotype). 

Robust lianas; stems twisted, costate, hollow, glabrous, internodes to 20 cm or more. 
Leaves ovate, to 20 cm long and 16 cm wide, margins entire, apices acute, bases cordate, 
upper surfaces glabrous, 5-7-nerved from the base, lower surfaces pilose; petioles to 7 cm 
long, glabrous. Inflorescence paniculate, the heads in spikes; ultimate branchlets angled, 
puberulent. Heads ca. 7 mm long, sessile to shortly pedicellate; exterior bract narrowly 
ovate, ca. Vz the length of the involucre, glabrate, apices acute; involucral bracts elliptic to 
oblong, ca. 3.5 mm long, glabrate to sparingly puberulent, 3-5-nerved, apices rounded, 
bases somewhat expanded and flattened; corollas ca. 4 mm long, white, tube 2-2.5 mm 
long, slightly enlarged at the base, throat turbinate-subcylindric, 1-1.5 mm long, teeth 
narrowly triangular, ca. 0.5 mm long. Achenes ca. 3.5 mm long, somewhat glandular, 
black with white ribs; pappus bristles ca. 35, white, ca. 4 mm long, barbellate, slightly 
thickened at the tips. 

Mikania aquaria, endemic to Peru, is well marked by its large ovate leaves 
with cordate bases and costate, hollow stems. The stems of this species are 
reported to be water filled. 

HUANUCO. Vicinity of Tingo Maria, S of bridge over Rio Chinchao on 
Huanuco-Tingo Maria Road, Mathias & Taylor 5904 (F). JUNfN. Near La 
Merced, Soukup 2509 (US). LORETO. Boqueron del Padre Abad, along Rio 
Chino, 300 m, Woytkowski 34388 (F). 

3. Mikania areolata Holmes & McDaniel, Phytologia 37: 467. 1977. TYPE: Peru, 
Amazonas, Cerros Calla Calla, 45 km above Balsas, midway on road to 



MACBRIDE: FLORA OF PERU 9 

Leimebamba, 3,100 m, June 19, 1964, Hutchison & Wright 5754 (F, holotype; 
MO, isotype). 

Lianas; stems terete, costate-sulcate after drying, finely puberulent, especially at the 
nodes, internodes to 20 cm long, nodes slightly enlarged. Leaves ovate, ca. 8 cm long and 
6 cm wide, margins entire to somewhat obscurely and remotely serrate, apices acute, 
bases cordate, upper surfaces puberulent, palmately 5-7-nerved from the base, nerves 
puberulent, lower surfaces puberulent-pilose, especially on the nerves, lighter than 
above, all veins exserted, tertiary veins forming areolae; petioles ca. 4 cm long, puberu- 
lent, opposite petioles connected by. a lacerate, puberulent stipule-like enation. In- 
florescence corymbose, ca. 6 cm long and 10 cm wide, bracteal leaves similar to cauline, 
only smaller; branchlets puberulent, branching trichotomously, heads usually disposed 
in 3's; pedicels 2-5 mm long. Heads 12-13 mm long; exterior bracts obovate, 6-7 mm long, 
somewhat pilose, apices rounded, bases cuneately narrowed into a petiole-like structure, 
3-5-nerved, ciliolate, borne at the top of the pedicel; involucral bracts ca. 10 mm long, 
puberulent, densely so at the tips, greenish, appearing to have lavender tips, apices 
rounded; corollas ca. 6 mm long, pale greenish-yellow; tube ca. 3.5 mm long; throat 
campanulate, ca. 2 mm long; teeth deltoid, ca. 0.5 mm long. Achenes ca. 4.5 mm long, 
black, ribs white; pappus bristles 90-100, whitish, ca. 5 mm long, barbellate. 

Mikania areolata, known only from the type, superficially resembles M. cor- 
difolia in habit, but may be distinguished by its terete rather than hexagonal 
stems of the latter species. The heads are also considerably larger, 12-14 mm 
long. 

4. Mikania arthroclada B. L. Robinson, Contr. Gray Herb. 80: 34. 1928. TYPE: 
Peru, Cuzco, above Cosnipata, between Tambos Trescruces and Tambamayo, 
2,500-2,600 m, May 7, 1912, Weberbauer 6972 (F, holotype, GH, photo; G, 
isotype, not seen). 

Twining shrubs; stems terete, densely fulvous-tomentulose, internodes to 6 cm long, 
nodes conspicuous, the internodes of branching stems leading to the inflorescence very 
short, 0.5-0.9 cm long. Leaves ovate-oblong, to 2 cm long and 1.5 cm wide, apices acute, 
margins serrate to vaguely crenate, bases cuneate, upper surfaces glandular-pubescent, 
pinnately nerved with secondary nerves separating near the base, lower surfaces some- 
what glandular-pubescent; petioles 2-5 mm long, puberulent. Inflorescence paniculate; 
branchlets somewhat hexagonal, fulvous-tomentulose; pedicels 1-2 mm long, fulvous- 
tomentulose. Heads 4.5-5 mm long; exterior bracts linear, ca. Vz the length of the in- 
volucre, borne at the base of the pedicel; involucral bracts oblong, ca. 3.5 mm long, 
glandular, puberulent, margins ciliolate; corollas ca. 2.5 mm long, white, tube ca. 1 mm 
long, throat ca. 1 mm long, gradually expanded, teeth deltoid, ca. 0.5 mm long. Achenes 
ca. 2.5 mm long, brownish; pappus bristles ca. 45, white, ca. 3 mm long. 

Mikania arthroclada, known only from the type material, is readily identified 
by its small leaves with coarsely serrate to somewhat crenate margins, dense 
fulvous-tomentulose indument, short internodes, and very prominent nodes 
from which the leaves appear to be early deciduous (especially from the lower 
nodes). 

5. Mikania aschersonii Hieron., Bot. Jahrb. Syst. 28: 577. 1901. TYPE: Colom- 
bia, Cauca, western slope of the Andes of Popayan, Lehmann 5979 (B, 
holotype, destroyed; F, isotype). Figure 1. 

M. aschersonii var. indula B. L. Robinson, Contr. Gray Herb. 77: 47. 1926. TYPE: Peru, 
Junin, La Merced, Hacienda Schunke, August 27-September 1, 1923, Macbride 5777 
(F, holotype; GH, isotype). 

Twining vines; stems striate-sulcate, glabrate to villous (when young), internodes to 11 
cm or more. Leaves ovate-lanceolate, to ca. 10 cm long and 3.5 cm wide, margins entire, 
apices acuminate, bases cuneate, upper and lower leaf surfaces glabrous, veins exserted; 



10 



FIELDIANA: BOTANY 



= 2 nun 




FIG. 1. Mikania aschersonii Hieron. A, habit; B, head; C, flower. (After McDaniel & 
Rimachi 22032 [IBE].) 



petioles 0.5-2 cm long, glabrous. Inflorescence corymbose; branchlets terete, villous to 
glandular-pubescent; pedicels 1-2 mm long, glandular-pubescent. Heads ca. 8 mm long; 
exterior bracts elliptic to ovate, ca. 4 mm long, apices acute; involucral bracts oblong, 
5-6.5 mm long, glabrate to glandular-pubescent especially toward the acute apex, bases 
somewhat enlarged or swollen; corollas ca. 4.5 mm long, tube ca. 1 mm long, throat 
subcylindric ca. 2.9 mm long, teeth triangular, ca. 0.6 mm long. Achenes ca. 3 mm long, 
dark; pappus bristles 50-60, reddish, 4-4.5 mm long, slender, barbellate. 

Mikania aschersonii is known from Peru and Colombia. Characters that may 
be used to identify this species include the oblong involucral bracts with 
slightly enlarged bases, somewhat in the manner of M. hookeriana, and corollas 
with a very short tube, about 1 mm long, with a subcylindric throat. 

CUZCO. Quispicanchis: Entre Inambari y 15 Mil, 500-600 m, Vargas 16519 
(US), La Concepcion, ca. 10.5 km NE Hacienda Luisiana and Apurimac River, 
Dudley 10559 (MO, US). HUANUCO. SW slope of Rio Llula, Pichis watershed 
on the ascent of Cerros del Sira, Dudley 13102 (US). JUNfN. Chanchamayo 
Valley, 1,500 m, Schunke s. n. (F). SAN MARTfN. Near Moyobamba, 1,100 m, 
Klug 3679 (F, GH, MO, US), Mariscal Caceres: Quebrada Luis Salas 15 km NE 
Puerto Pizana, 350-370 m, Schunke V. 6649 (MO). DPTO. UNKNOWN: Yanano, 
6,000 ft, Macbride 3775 (F, GH, US), Tambo de Vaca, 1,300 ft, Macbride 4906 (F, 
GH), Huacachi, estacion near Muna, 6,500 ft, Macbride 4151 (F, GH, US). 



MACBRIDE: FLORA OF PERU 11 

6. Mikania banisteriae DC., Prodr. 5: 193. 1836. TYPE: Brasil, 1834, Poeppig 135 
(G-DC, holotype, not seen, IDC microfiche 800. 819: II. 3; F, isotype). 

M. lanuginosa DC., Prodr. 5: 201. 1836. TYPE: Brasil, Slo Paulo, n. miss. 425 (G-DC, 

holotype, not seen, IDC microfiche 800. 819: II. 8). 
M. caudata Benth., PI. Hartw. 201. 1845. TYPE: Colombia, in sylvis prope Guaduas, 

Bogota, Hartweg 1112 (K, holotype, not seen). 
M. ruiziana Poepp. in Poepp. & Endl., Nov. Gen. Sp. PI. 3: 53. 1845. TYPE: Peru, 

Cuchero, Poeppig 1475 (W, holotype, not seen; B, not seen, F, isotypes). 
M. ruiziana var. lehmanniana Hieron., Bot. Jahrb. Syst. 19: 45. 1894. TYPE: Colombia, 

near Dolores, Tolima, 1,000-1,800 m, Lehmann 7483 (K, holotype, not seen; F, 

isotype). 
Willoughbya ferruginea Rusby, Mem. Torrey Bot. Club 6: 58. 1896. TYPE: Bolivia, 

Guanai-Tipuani, Bang 1419 (NY, holotype; F, US not seen, isotypes). 
W. hirsutissima Rusby, Mem. Torrey Bot. Club 6: 58. 1896, not W. hirsutissima Ktze., 

Rev. Gen. 1: 372. 1891. TYPE: not located. 
M. antioquiensis Hieron., Bot. Jahrb. Syst. 28: 580. 1901. TYPE: Colombia, prope Santa 

Domingo Cancan et Amalfi, 1,500-2,000 m, Lehmann 7971 (US, holotype; F, isotype). 
M. ferruginea (Rusby) Rusby ex Buchtien, Contr. Fl. Boliv. 1: 190. 1910. 
M. hirsutissima Buchtien, Contr. Fl. Boliv. 1: 190. 1910, notM. hirsutissima DC., Prodr. 5: 

200. 1836. TYPE: not seen. 
M. macbridei B. L. Robinson, Contr. Gray Herb. 73: 27. 1924. TYPE: Peru, Junin, La 

Merced, Hacienda Schunke, 4,000 ft, August 27-September 1, 1923, Macbride 5728 (F, 

holotype; GH, isotype). 
M. antioquiensis var. subcuneata B. L. Robinson, Contr. Gray Herb. 104: 32. 1934. TYPE: 

Peru, Loreto, Iquitos, trail to San Juan, Mexia 6492 (GH, holotype; US, F, NO, PH, 

isotypes). 
M. almagroi Cuatr. An. Univ. Madrid 4: 233. 1935. TYPE: Ecuador, San Jose, Isern 368 

(MA, holotype, not seen, F, fragm.). 
M. skutchii Blake, Brittonia 2: 333. 1937. TYPE: Costa Rica, El General, 1,130 m, Skutch 

2514 (US, holotype). 
M. canaquensis Badillo, Bol. Soc. Venez. Ci. Nat. 9: 132. 1944. TYPE: Venezuela, 

Canagua, Merida, Badillo 562 (VEN, holotype, not seen). 
M. bergantinensis Badillo, Bol. Soc. Venez. Ci. Nat. 10: 304. 1946. TYPE: Venezuela, 

Anzoatequi, Cerro Peonia, Steyermark 61540 (VEN, holotype, not seen; F, isotype). 
M. kavanaeyensis Badillo, I.e. 300. TYPE: Venezuela, Bolivar, Gran Sabana, between 

Mission of Santa Teresita de Kavanayen NW to Rio Karuai, Steyermark 59374 (VEN, 

holotype, not seen; F, isotype). 
M. meridana Badillo, I.e. 303. TYPE: Venezuela, Merida, Montana de San Jacinto, 

Steyermark 56667 (VEN, holotype, not seen; F, isotype). 
M. molinensis Badillo, I.e. 296. TYPE: Venezuela, Merida, near El Molino, between 

Canagua and Sta. Cruz de Mora, Badillo 610 (VEN, holotype, not seen; F, isotype). 
M. oniaensis Badillo, I.e. 297. TYPE: Venezuela, Merida, along Rio Onia, near Bolero, N 

of Mesa Bolivar, Steyermark 56736 (VEN, holotype, not seen). 
M. ptaretepuiensis Badillo, I.e. 302. TYPE: Venezuela, Bolivar, Ptaretepuf, Steyermark 

59936 (VEN, holotype, not seen; F, isotype). 

M. banisteriae DC. f. lanuginosa (DC.) Steyermark, Fieldiana, Bot. 28: 657. 1953. 
M. banisteriae f. lehmanniana (Hieron.) Steyermark, Fieldiana, Bot. 28: 657. 1953. 
M. banisteriae f. subglabra (B. L. Robinson) Steyermark, Fieldiana, Bot. 28: 657. 1953. 

Coarse lianas; stems terete, striate, normally villous to densely tomentose, rarely nearly 
glabrous, intemodes to 25 cm or more. Leaves narrowly to widely ovate, to ca. 20 cm long 
and 10 cm wide, apices acute to acuminate, bases cordate to truncate, margins entire to 
somewhat dentate, upper surfaces villous to tomentose, rarely glabrate, pinnately to 
subpalmately nerved, lower surfaces lighter than above, villous to tomentose; petioles to 
ca. 2 cm long, normally villous to tomentose. Inflorescence a large panicle; branchlets 
villous to tomentose to hirtellous; pedicels to ca. 4 mm long, densely pubescent. Heads 
7-9 mm long; exterior bracts ovate to obovate, ca. the same length as the involucre, 
glabrate to lightly pubescent with long, straight hairs, normally borne slightly below the 
involucre; involucral bracts ovate to oblong, 4-5 mm long, the outer pair pubescent with 
long hairs, the inner pair glabrate, or all nearly glabrate, apices rounded, the inner 



12 FIELDIANA: BOTANY 

surfaces generally pearly white; corollas 4.5-5 mm long, tube ca. 2x the length of the 
throat, throat broadly campanulate, teeth triangular, about the same length as the throat. 
Achenes 3-4 mm long; pappus bristles 40-50, white, 5-6 mm long, barbellate, often 
slightly thickened toward the tips. 

Mikania banisteriae is known from Mexico throughout Central America to and 
including northern South America to Peru, Bolivia, and Brazil. This species, as 
noted by the extensive synonymy, is highly variable and very widespread. A 
broad species concept, as applied by Steyermark (1953), is followed in this 
treatment. The primary differences in the proposed species (synonyms) involve 
leaf shape and kind and amount of pubescence, which, as Steyermark has 
shown, may vary on the same plant, and thus are totally unreliable for species 
delimitation. Reliable characters that can be used to identify this species, re- 
gardless of leaf shape and pubescence, include the large paniculate, rather 
dense inflorescence. 

The plants designated as Mikania rugosa, M. bullata, and M. trachodes by 
Robinson (1920, 1924), although very nearM. banisteriae, are herein regarded as 
distinct. For further information see those species in this work or Holmes & 
McDaniel (1979). Worthy of special mention is M. macbridei, which is, according 
to Robinson (1924), a well-marked species. It appears, upon examination of the 
type, that this name is based upon a specimen of M. banisteriae with all but the 
younger leaves covered with a fungal mycelium. It is identical in all other 
respects with that species. For further amplification see Holmes & McDaniel 
(1979). 

AMAZONAS. Bongara: 2-4 km WSW of Pomacocha, 2,200-2,400 m, Wurdack 
834 (US). ANCASH. Pampayacu, Kanehira 250 (US). CUZCO. Convencion: El 
Dorado, 2,900 m, Vargas 3509 (F); Paucartambo: Hda. Patria, Marin 1723 (F, US), 
km 123-124, Kosnipata, 3,000 m, Vargas 15481 (US). HUANUCO. SW slope of 
Rio Llula, Pichis watershed, ascent of Cerros del Sira, 1,290 m, Wolfe 12360 (US). 
JUNfN. Colonia Perene, 680 m, Killip & Smith 25013 (US), 590 m, Hitchcock 22129 
(GH), La Merced, Hacienda Schunke, 4,000 ft, Macbride 5512 (F, GH, US), Pichis 
Trail, Porvenir, 1,500-1,900 m, Killip & Smith 25907 (GH, US), Pichis Trail, 
Eneras, 1,600-1,900 m, Killip & Smith 25626 (F, GH, US), 25774 (US), Quimiri, 
Chanchamayo Valley, 1,500 m, Schunke 26 (F, GH), 475, 476 (both F), San 
Ramon, 900-1,300 m, Killip & Smith 24768 (GH, US), 24894 (F, GH, US). 
LORETO. Coronel Portillo: Plantacion Azul cerca Divisoria, Ricoutt s. n. (US), 
Maynas: Dtto. Iquitos, Iquitos, ca. 100 m, Killip & Smith 27478 (F, GH, US), 
Iquitos, Williams 3632 (F, GH), 1455 (F, GH), Iquitos, 120 m, Williams 8094 (F, 
GH), Iquitos, Puerto Almendro, Revilla 1479 (AMAZ, MO), Mishuyacu near 
Iquitos, 100 m, Klug 878 (F, US), Camino a Rumococha, 100-120 m, Ferreyra 
3370, Rio Nanay below Bellavista, trail to Picuruyacu, 170 m, McDaniel 16284 
(IBE, MO, NATC), trail from Picuru (lower Rio Nanay) to Rio Mazan, McDaniel 
et al. 21454 (AMAZ, GA, IBE, MO, NATC, US, USM), Quistococha to San Juan, 
McDaniel & Persinos 10706 (IBE, MO), lower Rio Nanay, Williams 438 (F, GH), 
Dtto. Indiana, Rio Manati, below San Isabel, McDaniel et al. 21652 (AMAZ, F, 
IBE, MO, NATC, NY, RB, US, USM), Loreto: Dtto. Nauta, Quebrada de Nauta, 
Croat 17532 (IBE, MO), Requena: Rio Tepiche, Santa Elena, McDaniel & Marcos 
11253 (IBE, MO); Yurac between Divisoria and Pucallpa, 15-20 km from Yurac, 
Allard 22140 (F, US). PUNO. Sandia, 2,600-3,000 m, Weberbauer 755 (GH). SAN 
MARTIN. Tarapoto, 530-580 m, McDaniel 13733 (IBE, MO), Tarapoto, Spruce 



MACBRIDE: FLORA OF PERU 13 

4079 (F, GH), San Roque, 1,300-1,500 m, Williams 7497 (F), alto Rio Huallaga, 
360-900 m, Williams 6602 (F, GH), Zepelacio, near Moyobamba, 1,200-1,600 m, 
Klug 3427 (F, GH, MO, US). 

7. Mikania brachyphylla Hieron., Bot. Jahrb. Syst. 36: 472. 1905. TYPE: Peru, 
Cajamarca, near Tambillo (Chanta Cruz), 2,600-2,700 m, Jelski 703 (B, 
holotype, destroyed, GH, photo & fragm.). 

Slender herbaceous twiners; stems somewhat 4-angled, purplish, glabrate, intemodes 
generally 2 cm long, but sometimes to 10 cm long. Leaves triangular-ovate, ca. 10 mm long 
and 7 mm wide, margins entire to obscurely 2-3-toothed on each side, apices acute to 
rounded, bases sagittate-cordate, upper surfaces scabrid, palmately 3-5-nerved, lower 
surfaces appressed-hirtellous, glandular in appearance; petioles to 5 mm long, opposite 
petioles connate by a stipular ring, with 3-5 ciliated appendages on each side of stem. 
Inflorescence a corymb; branchlets angular-sulcate, puberulent; pedicels ca. 4 mm long, 
angular, puberulent. Heads ca. 9 mm long; exterior bracts ovate, to 2 mm long, puberu- 
lent, borne slightly beneath the involucre; involucral bracts oblong, ca. 5 mm long, pu- 
berulent, apices obtuse; corollas ca. 5 mm long, yellowish (after drying), tube ca. 2.5 mm 
long, throat campanulate, 2 mm long, teeth deltoid, ca. 0.5 mm long, glandular. Achenes 
ca. 3 mm long, light olivaceous in color; pappus bristles ca. 75, buff in age, ca. 5 mm long, 
finely scabrid, thinning toward the tips. 

Mikania brachyphylla, known only from the type, is easily recognizable by 
somewhat quadrangular, purplish stems, corymbose inflorescence, and leaves 
ca. 1 cm or less long with scabrid upper surfaces. 

8. Mikania brevifaucia Holmes & McDaniel, Phytologia 37: 468. 1977. TYPE: 
Peru, Cuzco, Convencion, Tanamanche to Quellomayo, 3,700 m, July 25, 
1944, Vargas 4447 (F, holotype). 

Twiners; stems terete, striate to weakly costate, villous, soon glabrate to puberulent, 
internodes to ca. 13 cm long, nodes with stipule-like enations ca. 6 mm wide, laciniate 
with teeth 2-3 mm long. Leaves triangular, ca. 6 cm long and 3.5 cm wide, villous, 
glandular, apices acuminate, bases cordate, palmately 3-7-nerved, margins dentate to 
somewhat crenate, lighter below; petioles to ca. 2 cm long, villous; bracteal leaves similar 
to cauline, but reduced. Inflorescence corymbose, to ca. 9 cm wide and 6 cm long; 
branchlets terete, villous; pedicels 1-3 mm long, villous. Heads ca. 10 mm long; exterior 
bracts lanceolate to narrowly ovate, ca. 3.5 mm long, pilose to somewhat villous, apices 
acuminate; involucral bracts linear to lanceolate, densely pilose to villous, apices acumi- 
nate; corollas ca. 6 mm long, tube ca. 3.5 mm long, throat scarcely distinguishable from 
the tube, ca. 0.5 mm long, teeth narrowly triangular, ca. 2.5 mm long. Achenes 3-5 mm 
long, brownish with white ribs, slightly pubescent toward the top (near pappus bristles); 
pappus bristles ca. 70, white, ca. 5 mm long, weakly barbellate, narrowed toward the 
apex. 

Mikania brevifaucia is known only from Cuzco, Peru. Salient diagnostic 
characters include the triangular leaves with dentate to crenate margins and 
stipular appendages with lacinate teeth. 

CUZCO. Calca: Amparaes, borde de arroyo, 3,350 m, Chavez 3282 (MO). 

9. Mikania brooksii Holmes & McDaniel, Phytologia 41: 193. 1979. TYPE: Peru, 
Loreto, Maynas, Dtto. Iquitos, Rio Momon, trib. Rio Nanay, Santa Rosa, 
January 3, 1978, McDaniel, Rimachi & Brooks 21387 (IBE, holotype). 

Subligneous lianas; stems terete, somewhat puberulent-scabrid, sparsely glandular, 
intemodes to ca. 10 cm long. Leaves ovate to broadly ovate, to 8 cm long and 5 cm wide, 
bases obtuse to truncate, apices acute to acuminate, above somewhat scabrid, with 
whitish pustular hairs, the major nerves crisped-puberulent, pinnately nerved, with 2 



14 FIELDIANA: BOTANY 

pairs of secondary nerves separating within the basal l /3 of the leaf, the primary and 
secondary veins prominent, the others obscure, margins entire, below lighter, puberu- 
lent, all veins exserted; petioles 5-6 mm long, tomentulose-puberulent, thickish. In- 
florescence a panicle of oppositely borne corymbs, ca. 11 cm long and 8 cm in diameter, 
each corymb ca. 2 cm long and 4 cm in diameter, branching dichotomously; branchlets 
terete, densely crisped-puberulent, glandular. Heads ca. 7 mm long, sessile to very 
shortly pedicellate, ternately disposed; exterior bracts ovate-obovate to broadly elliptic, 
2.5 mm long, somewhat petiolate, apices rounded, irregularly ciliate, the midvein prom- 
inent, l /2- l /3 the length of the involucre, sparingly glandular; involucral bracts elliptic- 
oblong, 4-4.5 mm long, apex rounded, somewhat pubescent toward apex and base, 3-5 
obscure but visible veins present; corollas violet, ca. 3.5 mm long, tube and throat in- 
distinguishable, teeth deltoid, ca. 0.3 mm long, glandular. Achenes (slightly immature) to 
ca. 3 mm long, greenish; pappus bristles 40-45, white, ca. 3.5 mm long, scabrid, thinner 
toward the tips. 

LORETO. Maynas: Dtto. Iquitos. Rio Momon (trib. Rio Nanay), Santa Rosa, 
110 m, lowland forest rarely inundated, flowers pink, Rimachi 4832 (AMAZ, F, 
IBE, MO, NATC, US, USM); Rio Itaya, trocha de Sanangal, disturbed forest of 
restinga, 110 m, flowers lavender, McDaniel & Rimachi 23286 (AMAZ, F, FSU, 
IBE, MO, NATC, RB, TEX, US, USM). 

10. Mikania bulbisetifera Cuatr., Ann. Missouri Bot. Card. 52: 308. 1965. 
TYPE: Peru, Amazonas, Bangara, Jalca zone, along Yambrasbamba- 
Pomacocha trail between Yanayacu and Pomacocha, 2,300-2,400 m, June 26, 
1962, Wurdack 1078 (US, holotype; F, isorype). 

Vines; stems terete, costate, lanate, intemodes to ca. 3.5 cm long. Leaves lanceolate, 
coriaceous, ca. 7 cm long and 2 cm wide, margins subentire, apices acuminate, bases 
obtuse, upper surfaces scabrid when young, becoming somewhat muricate in age by loss 
of upper portion of hairs, lower surfaces lanose, lighter than above; petioles to 1.5 cm 
long, lanose. Inflorescence corymbo-paniculate, to ca. 8 cm long and 8 cm wide; branch- 
lets terete, lanose; pedicels 2-5 mm long, lanose. Heads 9-10 mm long; exterior bracts 
narrowly ovate, ca. % the length of the involucre, glabrate to sparingly pilose, somewhat 
irregularly ciliate, apices acute, pilose; involucral bracts ovate-oblong, 5-6 mm long, 
lightly pilose, apices acute to rounded, tomentulose; corollas ca. 4.5 mm long, white, tube 
ca. 1.5 mm long, throat campanulate, ca. 1.5 mm long, teeth narrowly triangular to lan- 
ceolate, ca. 1.5 mm long. Achenes glabrate to sparsely puberulent, ca. 3.2 mm long, 
grayish-brown, the ribs lighter in color; pappus bristles ca. 30, white, 5-6 mm long, 
barbellate, thickened at tips. 

Mikania bulbisetifera is known only from the type. The lanose pubescence and 
characters of the heads and inflorescence of this species suggest the highly 
variable M. banisteriae. However, the lanceolate leaves, the narrower exterior 
bract, and pappus bristles about one-half the number of M. banisteriae provide 
adequate distinction. 

11. Mikania bullata B. L. Robinson, Contr. Gray Herb. 61: 13. 1920. TYPE: 
Peru, Puno, between Tambo Ichubamba and Tambo Yuncacoyo, on the way 
from Sandia to Chunchusmayo, 1,800-2,000 m, June 7, 1920, Weberbauer 1085 
(B, holotype, destroyed, GH, photo & fragm.). 

Climbing shrubs; stems terete, densely covered with tawny spreading to deflexed 
soon-deciduous tomentum, internodes 5-13 cm long. Leaves ovate, to ca. 10 cm long and 
6 cm wide, margins obscurely denticulate, apices acute, bases cordate, upper surfaces 
chartaceo-coriaceous, tawny-pilose, bullate, scabrous, pinnately nerved, lower surfaces 
tawny-villous, punctate, veins exserted from the surface; petioles to ca. 2.5 cm long, 
tawny-tomenrulose, stout. Inflorescence a panicle, dense; branchlets tawny-tomentulose, 
leafy-bracted; pedicels 2-3 mm long, tawny-tomentulose. Heads 7-8 mm long; exterior 



MACBRIDE: FLORA OF PERU 15 

bracts linear to oblanceolate, ca. x /2 the length of the involucre, densely tawny-pilose, 
apices acute, bome at the summit of pedicel; involucral bracts oblong, 4-5 mm long, 
tawny-pilose, the outer darkened and somewhat punctate, apices rounded; corollas ca. 4 
mm long, tube ca. 1.5 mm long, throat broadly campanulate, ca. 1.25 mm long, teeth 
triangular, about as long as throat. Achenes 3-3.5 mm long, brownish-olivaceous; pappus 
bristles 35-40, white, ca. 5 mm long, barbellate, thickened at the tips. 

Mikania bullata was previously known only from the type. It is another 
species exceedingly near M. banisteriae. However, it differs in the scabrid na- 
ture of the upper leaf surfaces and the linear to oblanceolate exterior bract 
one-half or more as long as the involucre. Mikania banisteriae normally has 
smooth leaves and an ovate exterior bract about one-half as long as the in- 
volucre. The above description in part taken from Robinson (1922). 

PUNO. Sandia cerca a San Juan del Oro, valle Tampopata, 2,100-2,200 m, 
Ferreyra 16685 (MO, USM). 

12. Mikania carnosa Muschler, Bot. Jahrb. Syst. 50: 76. 1913. TYPE: Peru, 
Ayacucho, Huanta, Yanamonte, 2,700-3,000 m, June 1910, Weberbauer 5645 
(B, holotype, destroyed, GH, fragm.; F, isotype). 

Twining shrubs; stems terete to somewhat hexagonal, glabrate, internodes to 12 cm 
long. Leaves obovate to ovate to lanceolate-ovate, to ca. 4 cm long and 2 cm wide, margins 
entire near base, above serrate, the teeth callous-denticulate, apices rounded to somewhat 
acute, bases cuneate, upper surfaces glabrate, somewhat 3-5-nerved, lower surfaces pu- 
berulent; petioles to ca. 1 cm long. Inflorescence a corymb, 2-3 cm in diameter; branchlets 
angular-winged, glabrate; pedicels to ca. 5 mm long, glabrate. Heads ca. 10 mm long; 
exterior bracts ovate, ca. 4.5 mm long, glabrate, reticulate, apices acute to obtuse, borne at 
the top of the pedicel; involucral bracts lance-linear, ca. 8 mm long, glabrate, apices 
attenuate, somewhat irregularly ciliate; corollas ca. 6 mm long, pale yellow, tube ca. 3 mm 
long, throat ca. 1 mm long, teeth oblong, ca. 2 mm long. Achenes ca. 3.5 mm long, 
grayish; pappus bristles 75-80, yellow-white, ca. 6 mm long, scabrid. 

Mikania carnosa, known only from the type, may be readily distinguished by 
its lanceolate to ovate to obovate leaves with rounded apices and cuneate bases. 

13. Mikania chimborazensis Hieron., Bot. Jahrb. Syst. 29: 15. 1900. TYPE: 
Ecuador, forest at base of Mt. Chimborazo, Sodiro 767 (B, holotype, de- 
stroyed; F, isotype). 

Herbaceous to somewhat shrubby climbers; stems terete, striate, soon fistulose and 
glabrate, internodes to 15 cm long. Leaves ovate, ca. 10 cm long and 8 cm wide, margins 
serrate-crenate, apices acuminate, bases widely cordate, upper surfaces appressed- 
puberulent, scabrid, palmately 7-nerved from base, lower surfaces appressed puberulent; 
7-nerved; petioles ca. 5 cm long, somewhat puberulent, connected by a laciniate stipule- 
like ring. Inflorescence a 3-parted corymb, ca. 15 cm long and 10 cm wide; branchlets 
angular, puberulent. Heads 16-18 mm long; exterior bracts lance-elliptic, ca. 6-7 mm 
long, lightly puberulent, often purple-tinged; involucral bracts oblong, ca. 9 mm long, 
puberulent, purple-tinged, apices rounded, ciliate; corollas 9-11 mm long, purplish, tube 
and throat about the same length, throat turbinate, teeth triangular, ca. 1.5 mm long. 
Achenes 5-6 mm long, gray-olivaceous; pappus bristles ca. 75 or more, white, often 
purple-tinged, 10-12 mm long, barbellate. 

Mikania chimborazensis of Ecuador, Colombia, and Cajamarca, Peru has heads 
16-18 mm long, the largest of any known species of Mikania in Peru. 

CAJAMARCA. Celendin: canyon of Rio Maranon above Balsas, 2,930 m, 
Hutchison & Wright 5320 (US). 



16 FIELDIANA: BOTANY 

14. Mikania congesta DC., Prodr. 5: 197. 1836. TYPE: Puerto Rico, Bertero s.n. 
(G-DC, holotype, not seen, IDC microfiche 800. 820: II. 6). 

M. sieberiana DC., Prodr. 5: 196. 1836. TYPE: Trinidad, Sieber225 (G-DC, holotype, not 

seen, IDC microfiche 800. 820: II. 2, F, photo). 
M. parkeriana DC., I.e. 199. TYPE: British Guiana, Parker s.n. (G-DC, holotype, not 

seen, IDC microfiche 800. 821: III.4). 
M. variabilis Gardner, Lond. Jour. Bot. 5: 486. 1846. TYPE: Brazil, Alagoas, Piassabisu, 

Gardner 1344 (K, holotype, not seen). 
M. atriplicifolia Schultz-Bip. ex Miq., Stirp. Surin. Sel. 189. 1850. TYPE: In sylva prope 

Vredenburgerkreek, Focke s.n. (not seen). 

M. scandens (L.) Willd. var. congesta (DC.) Baker in Mart., Fl. Bras. 6: 249. 1876. 
M. scandens (L.) Willd. f. congesta (DC.) B. L. Robinson, Contr. Gray Herb. 64: 43. 1922. 

Twining or creeping lianas; stems striate to sulcate (after drying) to angled, puberulent 
to pilose, internodes 5-20 cm. Leaves deltoid-ovate to ovate-oblong, 2.5-14 cm long and 
2-10 cm wide, margins entire to weakly dentate to undulate, apices acute to attenuate, 
bases cordate, upper surfaces glabrous to puberulent, 3-5(7)-nerved from the base, lower 
surfaces puberulent; petioles 1.5-14 cm long, glabrous to puberulent. Inflorescence a 
dense corymbose-subglobose glomerule, ca. 1.5 cm in diameter; branchlets angled, pu- 
berulent. Heads 4-5 mm long, sessile to very shortly pedicellate; exterior bracts linear to 
lanceolate, as long as or exceeding the length of the involucre, pilose, apices attenuate; 
involucral bracts oval to oblong, 3-4 mm long, weakly pilose on the dorsal surface, apices 
mucronate; corollas ca. 3 mm long, white, glandular, tube ca. 1.5 mm long, throat ca. 1 
mm long, teeth deltoid, ca. 0.5 mm long. Achenes 1.5-2 mm long, glandular, black; 
pappus bristles 33-37, white, ca. 3 mm long, barbellate. 

Mikania congesta occurs on riverbanks, in swamps, or in other frequently 
inundated areas, to ca. 300 m elevation from Puerto Rico, the Lesser Antilles, 
and northern South America to Peru, Bolivia, and Brazil. This species is often 
confused with or treated as a form or variety of M. micrantha H.B.K. because of 
similar habit and foliage. The dense glomerule of M. congesta provides ample 
distinction from the open corymb of M. micrantha. 

LORETO. Middle Ucayali, Tessmann 3303 (NY); Maynas: Rio Ampiyacu, 
Pebas and vicinity, Plowman et al. 7064 (F), Caballo Cocha on the Amazon, 
Williams 2293 (F), Dtto. Indiana: Quebrada de Yanayacu, from mouth to "sac- 
arita," McDaniel et al. 21434 (AMAZ, F, IBE, MO, NATC, RB, US, USM); Dtto. 
Iquitos: Rio Itaya, Williams 120 (F), Rio Nanay near Iquitos, between Bellavista 
and Sta. Clotilde, McDaniel 13580 (FSU, IBE, MO), Rio Nanay, Morona Cocha, 
Rimachi 3608 (AMAZ, DUKE, F, FSU, GA, IBE, MO, NATC, NY, USM), Mis- 
huyacu, near Iquitos, Klug 933 (F, NY, US), Boca de Muyui, ca. 3 km W Iquitos, 
Croat 17494 (MO); Dtto. Mazan, Gamitanacocha, Rio Mazan, /. M. Schunke 54 
(F, NY, UC, US). MADRE DE DfOS. Cocha Cashu, Rio Manii, between Panagua 
& Tayakome, Foster 3461 (IBE). 

15. Mikania conglomerata Holmes & McDaniel, Phytologia 37: 469. 1977. 
TYPE: Peru, Huanuco, Pachitea, Honoria, Bosque Nacional de Iparia, a lo 
largo del Pachitea cerca del campamiento Miel de Abeja, October 24, 1967, 
Schunke V. 2249 (US, holotype; F, isotype). 

Lianas; stems terete, striate, with white, sessile glands, internodes to 30 cm long. 
Leaves broadly ovate, somewhat coriaceous, to ca. 22 cm long and 15 cm wide, margins 
entire, revolute, apices caudate, bases truncate to subcordate, upper surfaces glabrate, 
subpinnately 5-7-nerved from near the base, tertiary veins prominent, transverse, lower 
surfaces glabrate, veins exserted from the surface; petioles to 5 cm long, sulcate. In- 
florescence a panicle; heads sessile, arranged in a very dense subspherical glomerule to 
ca. 2 cm in diameter; branchlets angular-flattened, puberulent. Heads ca. 6 mm. long; 



MACBRIDE: FLORA OF PERU 17 

exterior bracts elliptic-oblong, ca. 1.5 mm long, puberulent; involucral bracts elliptic- 
oblong, ca. 4 mm long, puberulent, apices rounded, ciliolate; corollas ca. 4 mm long, 
white, scarcely fragrant, tube ca. 1.5 mm long, throat narrowly campanulate, ca. 2 mm 
long, teeth deltoid, ca. 0.5 mm long. Achenes ca. 2 mm long (immature); pappus bristles 
ca. 35, white, ca. 4 mm long, barbellate, slightly thickened at the tips. 

Mikania conglomerate!, known only from the type, is characterized by its large 
(up to 22 cm long), somewhat coriaceous leaves with caudate apices. 

16. Mikania cordifolia (L. f.) Willd., Sp. PI. 3: 1746. 1803. 

Cacalia cordifolia L. f., Suppl. PI. 351. 1781. TYPE: Colombia, Mutis 1818 (LINN, 

holorype not seen, IDC microfiche 117. 555: 1.7; US, isotype). 
Mikania mollis H.B.K., Nov. Gen. Sp. PI. 4: 105. ed. fol. 1818. TYPE: Peru, 

Guanacabamba River, Humboldt & Bonpland (P, holorype). 
M. suaveolens H.B.K., Nov. Gen. Sp. PI. 4: 106. ed. fol. 1818. TYPE: Colombia, near 

Guaruma, Humboldt & Bonpland (P, holorype). 
M. poeppigii Spreng., Syst. Veg. 3: 422. 1826. TYPE: Cuba, Poeppig (P, holotype, not 

seen; W, isotype, not seen). 
M. cissampelina DC., Prodr. 5: 195. 1836. TYPE: Santo Domingo (G-DC, holotype, not 

seen). 
M. gonoclada DC., Prodr. 5: 199. 1836. TYPE: Mexico, Tampico, Tamaulipas, Berlandier 

137 (G-DC, holotype, not seen, IDC microfiche 800. 821: II. 4). 
M. convolvulacea DC., Prodr. 5: 199. 1836. TYPE: Santo Domingo (G-DC, holotype, not 

seen, IDC microfiche 800. 819: III. 7). (See Robinson, Contr. Gray Herb. 104: 70. 1934.) 
M. surinamensis Miq., Stirp. Sur. Select. 188. 1850. TYPE: Surinam, prope plantionem 

La Rencontre, Focke (U, holotype, not seen). 
M. hostmanii Miq., Stirp. Sur. Select. 189. 1850. TYPE: Surinam, Hostmann 717 (K, 

holotype). 
M. scandens (L.) Willd. var. rhodotricha Baker in Mart., Fl. Bras. 6: 250. 1876. TYPE: 

Brazil, Rio de Janeiro, Serra dos Orgaos, Gardner 60 (K, holotype). 
Eupatorium marquesianum Gomez de la Maza, Dice. Bot. Nom. Vulgares Cubanos & 

Puerto-Riquenos 54. 1889. TYPE: not seen. 
Willoughbya cordifolia (L.f.) Kuntze, Rev. Gen. PI. 1: 372. 1891. 
Mikania vellosiana B. Rodr., Hort. Flum. 276. 1895. TYPE: not seen. 
Willoughbaeya halei Small, Fl. S.E.U.S. 1170. 1903. TYPE: United States, Louisiana, Hale 

(GH, holotype?). 
Mikania yapasensis B. L. Robinson, Contr. Gray Herb. 104: 48. 1934. TYPE: Peru, Junin, 

Pichis Trail, Yapas, 1,350-1,600 m, June 28-29, 1929, Killip & Smith 25493 (GH, 

holotype). 

Herbaceous to semiwoody twiners; stems hexagonal, gray-tomentulose or tomentose, 
usually spirally twisted, internodes 5-20 cm long. Leaves ovate, 5-10 cm long and 3-8 cm 
wide, margins subentire to undulate-dentate, apices acute to acuminate, bases cordate 
with a wide and open sinus, upper surfaces densely pilose to tomentose, pale green, 
3-5-nerved from the base, lower surfaces densely pilose to tomentose, lighter than above; 
petioles 2-5.5 cm long, veins appearing slightly decurrent upon petiole, densely pilose to 
tomentose, opposite petioles connected by a transverse ridge with a small laciniate 
stipule-like appendage. Inflorescence a compound corymb, terminal and lateral, ca. 6cm 
long and 7 cm wide; branchlets hexagonal, densely pilose to tomentose; pedicels 0.5-3 
mm long, hexagonal, pilose to tomentose. Heads 7-10 mm long; exterior bracts linear to 
narrowly lanceolate, 4-5 mm long, densely pilose, apices acuminate; involucral bracts 
elliptic to narrowly ovate, 6-8 mm long, substramineous, apices acute to slightly rounded; 
corollas 3.5-5 mm long, white, tube 1.5-2 mm long, throat 1-1.5 mm long, glabrous, teeth 
linear, slightly exceeding the length of the throat, recurved when dry. Achenes 3-4 mm 
long, glabrous to pubescent, lightly glandular, brown; pappus bristles white, ca. 60, 
thinning toward the tips, barbellate. 

Mikania cordifolia is a very widespread species occurring from Gulf coastal 
United States (Louisiana, Mississippi, and Florida) throughout the wet tropical 
Americas to Argentina. This species is easily recognized by the prominently 



18 FIELDIANA: BOTANY 

hexagonal stems normally tomentulose to tomentose, the compound corymbose 
inflorescence, and corolla teeth about the same length as or slightly longer than 
the throat, the teeth usually reflexed or crisped in the dried state. 

AYACUCHO. Aina, between Huanta and Rio Apurimac, 750-1,000 m, Killip 
& Smith 22736 (US). CUZCO. Convention, valley of Rio Vilcanota at mouth of 
Rio Sambray, 1,000 m, Mexia 8042 (MO, NY, TEX, UC, US). HUANUCO. 
Huanuco, Tingo Maria, valley of Rio Huallaga, trocha a la Cueva de las Pavas, 
700-800 m, Ferreyra 4244 (MO, US, USM), Rio Huallaga Canon, below Rio Santo 
Domingo, ca. 4,000 ft, Macbride 4199 (US). JUNfN; Quempiri, Rio Ene, 46CM80 
m, Ferreyra 26381 (US), Colonia Perene, 680 m, Killip & Smith 24991 (US) and 
25239 (US), Tarma, entre La Merced y Quimiri, valle de Canchamayo, 800-900 
m, Ferreyra 3682 (US). LORETO. Ucayali: Contamana, near road to Oriente, 160 
m, McDaniel 14041 (IBE). MADRE DE DfOS. Manu: Hacienda S. Antonio, Var- 
gas 14664 (US), Rio Acre, Ule 9884 (L). SAN MARTfN. Silcayo, cerca a Tarapoto, 
Ferreyra 7768 (US), Lamas: arriba de San Jose de Sisa, selva tropical, 500-550 m, 
Ferreyra 7917 (MO, USM). 

17. Mikania crassifolia Hieron., Bot. Jahrb. Syst. 36: 471. 1905. TYPE: Peru, 
Cajamarca, Tambillo, October 30, 1878, Jelski 696 (B, holotype, destroyed; 
MO, US, isotypes). 

Somewhat shrubby twiners; stems terete, papillate-scabrid, internodes to 30 cm or 
more long. Leaves ovate, to ca. 14 cm long and 9 cm wide, slightly fleshy, margins entire 
or with a few inconspicuous teeth, apices acuminate, bases rounded, somewhat cuneate 
upon the petiole at the point of insertion, subpinnately 5-7-nerved from near the base; 
petioles to ca. 4 cm long, opposite petioles connected by stipule-like enations of ca. 1.5 cm 
long and 0.6 cm wide, subentire to dentate. Inflorescence a rather dense corymb, to ca. 3 
cm in diameter; pedicels 2-3 mm long, puberulent. Heads ca. 13 cm long; exterior bracts 
lance-linear, 6-8 mm long, apices acute, borne at the top of pedicel; involucral bracts 
oblong, 8-9 mm long, glandular, apex narrowed to an obtuse point, somewhat ciliolate 
and purplish in color, prominently nervate; corollas ca. 8 mm long, dark violet, tube 3-4 
mm long, gradually enlarged into a funnelform throat, throat 2 mm long, teeth triangular, 
ca. 2 mm long. Achenes ca. 5.5 mm long, olivaceous, hispid on the angles; pappus bristles 
ca. 120 or more, white, ca. 8 mm long, barbellate. 

Mikania crassifolia, known only from the type, is characterized by its some- 
what fleshy leaves, heads ca. 13 mm long, and corolla reported to be deep 
purple in color. 

18. Mikania cutervensis Hieron., Bot. Jahrb. Syst. 36: 473. 1905. TYPE: Peru, 
between Chota and Cutervo, June 1879, Jelski 795 (B, holotype, destroyed). 
Kanimia cutervensis (Hieron.) Hieron., Bot. Jahrb. Syst. 40: 394. 1908. 

Suffrutescent (?) twining lianas; stems subangular, sparingly villose, at length terete, 
glabrous, greenish, internodes to ca. 16 cm long. Leaves subtriangular to cordate-ovate, 6 
cm long and 5 cm wide, apices acute-acuminate, margins coarsely dentate, the teeth 
semisubcircular-triangular, mucronate, ca. 4 mm long and 3-7 mm apart, above glabrate, 
the nerves sparingly pubescent, 5-7-nerved from the base, below villous, especially on 
the nerves; petioles to 4 cm long, often provided at the base with subulate, stipule-like 
enations ca. 2 mm long. Inflorescence corymbose. Heads ca. 12 mm or more long, sessile 
to subsessile; exterior bracts lanceolate, acute, glabrous, ca. 7 mm long, 3-nervate, 
greenish- to whitish-purple, margins ciliate; involucral bracts lanceolate, ca. 12 mm long, 
acute, scarious, greenish-stramineous, 3-nervate; corollas purple, ca. 8 mm long, gla- 
brous, the tube ca. 1.5 mm long, throat infundibuliform, ca. 4.5 mm long, teeth lanceo- 
late, ca. 2 mm long. Achenes (immature) ca. 5 mm long, glabrous; pappus bristles ca. 
100-105, reddish, ca. 8.5 mm long, apices not thickened. 



MACBRIDE: FLORA OF PERU 19 

The above description was drawn from that of Hieronymus. The species 
appears to be characterized by its more or less triangular leaves with coarsely 
dentate margins. 

The specimen listed below, although very immature, is cited with this species 
because it appears to agree in essentials. 

HUANUCO. Cajan, Huallaga, Weberbauer 4036 (F, fragm.). 



19. Mikania cuzcoensis Holmes & McDaniel, Phytologia 35: 329. 1977. TYPE: 
Peru, Cuzco, Quispicanchis, entre Inambari y 15 Mil, 500-650 m, September 3, 
1965, Vargas 16521 (US, holotype). 

Somewhat shrubby or procumbent lianas; stems terete, costate (after drying), glabrous, 
internodes to ca. 20 cm long. Leaves ovate, 6-10 cm long and 5-8 cm wide, margins entire, 
apices acute to attenuate, bases cordate, upper surfaces glabrous, 5-7-nerved from the 
base, pale green, lower surfaces with dark globular sessile glands, lighter than above; 
petioles 3-8 cm long, thin, puberulent. Inflorescence a racemosely disposed panicle, ca. 15 
cm long and 15 cm wide; branchlets slightly angular, puberulent; pedicels ca. 5 mm long, 
puberulent. Heads 5-6 mm long; exterior bracts linear, 2-2.5 mm long, borne at the 
summit of the pedicel; involucral bracts oblong, ca. 4 mm long, glabrate, apices obtuse, 
bases slightly narrowed, the center portion of each of the inner bracts extended into a 
small knoblike protuberance; corollas funnelform to slightly campanulate, ca. 2.5-3 mm 
long, tube 1.2-1.5 mm long, throat 0.5-0.8 mm long, teeth triangular, ca. 0.7 mm long. 
Achenes ca. 2.5 mm long, glabrous, dark brown; pappus bristles 35-40, white, ca. 4 mm 
long, barbellate, slightly thickened toward the tips. 

Mikania cuzcoensis, endemic to Peru, is characterized by the large racemosely 
disposed panicle with heads loosely arranged on pedicels ca. 5 mm long. 

CUZCO. Quispicanchis, entre Mashiche y Tuanssan, 510 m, Vargas 16450 
(US). HUANUCO. Tulumayo, cerca Tingo Maria, Carretera Huanuco-Pucallpa, 
700-800 m, Ferreyra 2159 (US, USM). 



20. Mikania decora Poepp. in Poepp. & Endl., Nov. Gen. Sp. PI. 3: 53. 1845. 
TYPE: Peru, Ancash, Pampayaco, woods around Cuchero, Poeppig 1645 (W, 
holotype, F, photo). Figure 2. 

Willoughbya bangii Rusby, Mem. Torrey Bot. Club 3: 53. 1893. TYPE: Bolivia, La Paz, 

Yungas, Bang 523 (NY, holotype; GH, isotype). 
Mikania tarapotensis B. L. Robinson, Contr. Gray Herb. 61: 21. 1920. TYPE: Peru, San 

Martin, mountains along River Mayo, near Tarapoto, Spruce 4822 (GH, holotype, F, 

photo & fragm.). 
M. decora var. heteroneura B. L. Robinson, Contr. Gray Herb. 73: 23. 1924. TYPE: Peru, 

Villacabamba, hacienda on Rio Chinchao, 610 m, July 17-26, 1923, Macbride 5167 (F, 

holotype; GH, isotype). 

Twining lianas or shrubs; stems terete, costate, fistulose, at first puberulent, soon 
glabrate, internodes to 12 cm long. Leaves ovate-oblong, 6-15 cm long and 7 cm wide, 
margins entire, apices acuminate, bases rounded, upper surfaces glabrous, glandular, 
5-7-nerved from near the base, lower surfaces puberulent, glandular, nerves exserted; 
petioles ca. 1-1. 5 cm long, puberulent. Inflorescence paniculate, lax, ultimate branchlets 
disposed in racemes; branchlets puberulent; pedicels 3-5 mm long, filiform, puberulent. 
Heads 4-5 mm long; exterior bracts subulate, ca. 1.5 mm long, puberulent, borne at the 
base of the pedicel; involucral bracts oblong, 3-3.5 mm long, glabrate, striate, apices 
obtuse, puberulent; corollas 2.4-2.5 mm long, whitish green, tube ca. 1 mm long, glan- 
dular, throat semicampanulate, ca. 1 mm long, teeth triangular, ca. 0.5 mm long. Achenes 
ca. 1.5 mm long, brownish; pappus bristles ca. 40, pinkish white, ca. 3 mm long, barbel- 
late. 




= 2 mm 



FIG. 2. Mikania decora Poepp. A, habit; B, head; C, flower. (After McDaniel & Rimachi 
22033 [IBE].) 



20 



MACBRIDE: FLORA OF PERU 21 

Mikania decora occurs to ca. 3,000 m elevation in Peru and Bolivia. The lax 
racemose inflorescence with heads borne on slender pedicels 3-5 mm long 
characterizes this species. 

AMAZONAS. Bongara: 2-4 km WSW of Pomacocha, 2,200-2,400 m, Wurdack 
839 (US). CAJAMARCA. Jaen: Valley of the River Tabaconas, 1,200 m, Weber- 
bauer 6254 (F, GH, US). CUZCO. Villacabamba: Hacienda on Rio Chinchao, 
Macbride 5191 (F, GH). HUANUCO. Carpish, carretera Huanuco-Tingo Maria, 
2,800-2,900 m, Ferreyra 2393 (MO, US, USM). JUNfN. Schunke Hacienda, above 
San Ramon, 1,300-1,700 m, C. Schunke A2 (US). SAN MARTfN. Tarapoto, car- 
retera Tarapoto-Yurimaguas, km 12-15, ca. 2,250 ft, McDaniel & Rimachi 22033 
(AMAZ, F, FSU, GA, IBE, MO, NATC, NY, US, USM), Zepelacio, near 
Moyobamba, 1,200-1,600 m, Klug 3531 (F, GH, MO, US), 3533 (F, GH, MO), and 
3655 (GH, MO). 

21. Mikania desmocephala B. L. Robinson, Contr. Gray Herb. 64: 7. 1922. 
TYPE: Bolivia, near Yungas, 1,220 m, Rusby 1740 (NY, holotype). 

M. desmocephala var. peruviana B. L. Robinson, Contr. Gray Herb. 77:50. 1926. TYPE: 
Peru, Junin, La Merced, Hacienda Schunke, 1,220 m, Macbride 5776 (F, holotype). 

Twining vines; stems terete, costate, densely pinkish-puberulent, internodes to ca. 12 
cm long. Leaves narrowly ovate, to ca. 11 cm long and 5 cm wide, margins entire, rev- 
olute, apices attenuate, bases rounded, upper surfaces lightly scabrid, 5-nerved from near 
the base, lower surfaces glabrate, glandular; petioles 2-3 cm long, scabrid. Inflorescence 
paniculate, heads disposed in dense hemispherical glomerules ca. 1.5 cm in diameter; 
branchlets densely puberulent. Heads sessile, 7-8 mm long; exterior bracts obovate, ca. 
Va-Vz the length of the involucre, puberulent, apices acute; involucral bracts oblong, ca. 
5.5 mm long, sparsely pilose, apices rounded, bases slightly swollen; corollas ca. 4 mm 
long, tube ca. 2 mm long, throat campanulate, 1.3-1.5 mm long, teeth deltoid, less than Vz 
the length of the throat. Achenes ca. 3 mm long, black, ribs white; pappus bristles 40-45, 
white, 3.5-4 mm long barbellate. 

Mikania desmocephala of Peru and Bolivia may be distinguished from the 
similar M. hookeriana by its narrower leaves with revolute margins. 

CUZCO. La Convencion, almost to Camp 3 from Camp 2 l /2, ca. 1,700 m, 
Dudley 10446 (MO, US, USM). JUNfN. Pichis Trail, Dos de Mayo, 1,700-1,900 
m, Killip & Smith 25800 & 25824 (both at GH, US). LORETO. Coronel Portillo: 
Divisoria, entre Tingo Maria y Pucallpa, 1,600-1,700 m, Ferreyra 4157 (MO, US, 
USM). 

22. Mikania dudleyi Holmes & McDaniel, Phytologia 45: 297. 1980. TYPE: 
Peru, Cuzco, La Convencion, exposed steep ridge at top of old landslide 
below Camp 5, ca. 2,800 m, July 9, 1968, Dudley 10913 (US, holotype). 

Subligneous lianas; stems terete, costate-striate, glabrate, internodes to 18 cm long. 
Leaves ovate-oblong, coriaceous, dark green, glossy, to 13 cm long and 6 cm wide, mar- 
gins entire to remotely and obscurely denticulate, apices attenuate, bases narrowly cor- 
date (bracteal leaves subcordate to truncate), upper surfaces glabrate to puberulent, glan- 
dular, subpinnately nerved from near the base, with 2 pairs of secondary nerves separat- 
ing within 1 cm of the base thence arching toward the apex, tertiary veins exserted; 
petioles to 2.5 cm long, thickish. Inflorescence a panicle, to ca. 12 cm long and 7 cm in 
diameter; branchlets terete, crisped-puberulent; pedicels ca. 1 mm long, terete. Heads 5-6 
mm long; exterior bracts linear-lanceolate, ca. 1.5 mm long, puberulent, borne at the base 
of the pedicel; involucral bracts linear to elliptic-oblong, ca. 4 mm long, puberulent, 
apices somewhat acute, densely puberulent-pilose; corollas violet, ca. 3.5 mm long, glan- 
dular, tube ca. 1.3 mm long, throat campanulate, ca. 1.6 long, teeth broadly triangular, ca. 



FIELDIANA: BOTANY 

0.6 mm long, glandular, puberulent. Achenes ca. 1.5 mm long (immature), brownish; 
pappus bristles ca. 30, white, ca. 4.5 mm long, barbellate, thickened at the tips. 

Mikania dudleyi, known only from the type, is characterized by its ovate 
leaves with narrowly cordate bases. 

23. Mikania ecu ado re ns is Holmes & McDaniel, Phytologia 31: 274. 1975. TYPE: 
Ecuador, Guayas, Camp -3563 (MICH, holorype; BR, NY, RB, isotypes). 

Twining herbs; stems 4-6 angled (after drying), lightly to moderately villous with 
sordid-white, jointed hairs, intemodes 10-15 cm long. Leaves ovate, 2-5 cm long and 
1.5-4 cm wide, margins entire to crenate-undulate, apices acute to acuminate, bases 
cordate, the sinus wide, upper surfaces pilose, lower surfaces velutinous with sordid- 
white hairs, 3-7-nerved from the base; petioles 1-5 cm long, densely pubescent. In- 
florescence a rounded to flat-topped corymb, sometimes rather dense, 1-3 cm long and 
1-6 cm wide; branchlets with villous hairs; pedicels ca. 1-3 mm long. Heads 4-4.5 mm 
long; exterior bracts linear to lanceolate, ca. 2.5 mm long, apices slightly erose to acumi- 
nate, outer surfaces pilose; involucral bracts lanceolate, 3-4 mm long, apices acute to 
acuminate, pilose, obscurely 1-3-nerved; corollas 2-2.5 mm long, white with globular 
glands toward the summit, tube ca. 1-1.3 mm long, gradually expanding into the throat, 
teeth broadly triangular, ca. 0.5 mm long. Achenes ca. 1.5 mm long, glandular, brownish; 
pappus bristles ca. 22, 1.5 mm long, white, thick, nonbarbellate. 

Mikania ecuadorensis has been found only in the lower coastal plain of Ecuador 
and nearby Peru. It has affinities with both M. micrantha and M. congesta but 
can be distinguished by its thick pappus of about 22 bristles rather than a thin, 
barbellate pappus of about 35 bristles in the other two species. 

PIURA. Piura, Townsend 791 (F); Talara, Haught 83 (GH, US). 

24. Mikania featherstonei B. L. Robinson, Contr. Gray Herb. 68: 38. 1923. 
TYPE: Peru, Huanuco, Pueblo, 3 mi below Ambo, September 19, 1922, Mac- 
bride cV Featherstone 2428 (F, holotype). 

Twining vines; stems terete, older stems sulcate, rusty puberulent, internodes to ca. 11 
cm long. Leaves ovate, somewhat coriaceous, to ca. 14 cm long and 9 cm wide, margins 
entire to irregularly and coarsely serrate to revolute, apices acute to acuminate, bases 
cordate, bracteal leaves with obtuse to truncate to somewhat cuneate bases, upper sur- 
faces rugose, scabrid, subpinnately to pinnately 5-7-nerved, lower surfaces hispid to 
somewhat scabrid, nerves prominent; petioles to ca. 4 cm long, hispid to scabrid, sulcate. 
Inflorescence a corymb, to ca. 5 cm long and 10 cm wide; branchlets angled to flattened, 
rusty hispid to scabrid. Heads 12-13 mm long, sessile to shortly pedicellate (to ca. 3 mm); 
exterior bracts linear, to ca. 4 mm long, somewhat hispid, borne at base of head; in- 
volucral bracts elliptic-oblong, 8-9 mm long, outer pair hispid to scabrid intermixed with 
puberulent hairs, the inner glabrate to lightly puberulent, the apices rounded, puberu- 
lent; corollas salverform, ca. 6 mm long, greenish- white, tube ca. 4.5 mm long, throat ca. 
0.2 mm long, teeth deltoid, ca. 1.3 mm long; anthers purple. Achenes ca. 5 mm long, 
black; pappus bristles ca. 75, cameous, ca. 6 mm long, barbellate. 

Mikania featherstonei is a well-marked species known only from the type. It 
may be distinguished by its rusty puberulent stems and ovate, pinnately veined 
leaves with scabrid upper surfaces. 

25. Mikania filicifolia B. L. Robinson, Contr. Gray Herb. 61: 14. 1920. TYPE: 
Peru, Cuzco, Machu Picchu, ca. 2,100 m, Cook & Gilbert 853 (US, holotype). 

Trailing vines; stems sulcate-costate, purplish-brown, puberulent at first, soon gla- 
brate, intemodes to ca. 10 cm long. Leaves 2-3-pinnately divided, 3-4 cm long and 2-3 cm 
wide, leaflets ovate-orbicular, ca. 5 mm in diameter, entire, obtuse at apex, terminal 
leaflet lanceolate, attenuate at tip, petiolulate, surfaces glabrate to puberulent; petiole to 



MACBRIDE: FLORA OF PERU 23 

ca. 1.5cm long, puberulent. Inflorescence a trifid corymb, ca. 7cm in diameter; branchlets 
somewhat angular, puberulent; pedicels 1-5 mm long, puberulent. Heads 8-9 mm long; 
exterior bracts narrowly ovate to obovate, ca. Vz the length of the involucre, glabrate, 
apices acute-attenuate, borne slightly beneath the head; involucral bracts lance-linear, ca. 
7 mm long, glabrate, apices attenuate, bases slightly tumid; corollas ca. 5 mm long, 
glandular, tube ca. 2.5 mm long, throat ca. 1.0 mm long, teeth linear, ca. 1.5 mm long. 
Achenes tan, ca. 2.5 mm long, lightly glandular at the summit, ribs white; pappus bristles 
ca. 40, white, 4-5 mm long, finely scabrid. 

Mikania filicifolia is a very striking species known only from the type. The 
delicate, pinnately dissected, femlike foliage easily separates this species from 
the only other species of this genus in Peru known to have divided leaves, M. 
ternata, which has palmately divided leaves. 

26. Mikania glandulifera Holmes & McDaniel, Phytologia 37: 470. 1977. TYPE: 
Peru, Huanuco, Pachitea, Honoria, Bosque Nacional de Iparia, Rio Pachitea 
ca. del campamento Miel de Abeja, January 6, 1967, Schunke V. 1487 (US, 
holotype; F, IBE, isotypes). 

Lianas; stems terete, striate-costate, glabrate, with dark glandular punctations (espe- 
cially near nodes), intemodes to ca. 16 cm long. Leaves elliptic-ovate, membraneous, to 
ca. 13 cm long and 6 cm wide, margins entire, apices attenuate-caudate, bases truncate to 
obtuse, upper surfaces glabrate, with dark glandular punctations near midvein, pinnately 
veined, lower surfaces with dark glandular punctations; petioles to ca. 3 cm long, with 
dark glands. Inflorescence a panicle, the ultimate segments spicately arranged; branchlets 
angular, puberulent, with somewhat stalked dark glands. Heads sessile, ca. 6 mm long; 
exterior bracts lance-elliptic, to ca. 1.5 mm long, apices acute, often with dark glands; 
involucral bracts elliptic-oblong, ca. 5 mm long, glabrate, with a few dark glands, apices 
rounded, densely pilose, bases somewhat swollen; corollas ca. 3 mm long, white, tube ca. 
1.5mm long, throat ca. 0.8mm long, turbinate, teeth lanceolate, ca. 0.7mm long. Achenes 
ca. 2 mm long, brownish; pappus bristles 35-40, white, ca. 3.5 mm long, scabrid. 

Mikania glandulifera is characterized by the strictly spicately disposed heads 
and elliptic-oblong membraneous leaves. 

27. Mikania guaco H. & B., PI. Aeq. 2: 84. 1809. TYPE: Colombia, Magdalena 
River between Mahates & Angostura, Humboldt & Bonpland 1447 (P, holotype, 
not seen). Figure 3. 

Mikania attenuata DC., Prodr. 5: 195. 1836. TYPE: Peru, "montibus Huanacocensis," 

Haenke (G-DC, holotype, not seen, IDC microfiche 800. 820: I.I, GH, photo). 
M. brachiata Poepp. in Poepp. & Endl., Nov. Gen. Sp. PI. 3: 53. 1845. TYPE: Peru, in 

forests at Tocache, Poeppig 2041 (W, holotype, not seen, F, photo & fragm.). 
M. cuneata Schultz-Bip., Bonplandia 4: 54. 1856. (nomen nudum). TYPE: Peru, San 

Govan, Lechler 2477 (F, isotype). 

M. amara var. guaco (H. & B.) Baker in Mart., Fl. Bras. 6: 237. 1876. 
Willoughbya guaco (H. & B.) Kuntze, Rev. Gen. 1: 372. 1891. 
M. olivacea Klatt, Bull. Soc. Roy. Bot. Belg. 31: 195. 1892 (1893). TYPE: Costa Rica, 

forests of Buenos Aires, 2,300 m, Pittier4433 (BR, holotype; GH, isotype). 

Lianas; stems terete, pithy to hollow, glabrous, internodes to 20 cm or more long. 
Leaves ovate-oval, thin, to 20 cm or more long and 12 cm or more wide, margins entire to 
somewhat denticulate, apices attenuate, bases acuminate and prominently decurrent 
upon the petiole, upper surfaces puberulent to sometimes scabrid, subpinnately 5-7 
(9)-nerved from near the base, lower surfaces glabrate; petioles to 5 cm long, thin. In- 
florescence a corymb, the ultimate segments rather dense, to ca. 6 cm long and 12 cm 
wide; branchlets angled, puberulent. Heads sessile to shortly pedicellate, disposed in 
groups of 3's, 8-10 mm long; exterior bracts subulate to linear, 0.5-2 mm long, puberu- 
lent; involucral bracts oblong, 4-6 mm long, puberulent, especially toward the rounded 
apices; corollas ca. 5 mm long, white to lilac to brownish, tube ca. 2 mm long, throat 2.4 



24 FIELDIANA: BOTANY 

mm long, teeth deltoid, ca. 0.6mm long, style branches prominently hirsute. Achenesca. 
3 mm long, blackish; pappus bristles ca. 60, buff (in age), 4-4.5 mm long, barbellate, 
thinner toward the tips. 

Mikania guaco is a very well-known and widespread species that occurs from 
southern Mexico throughout Central America to northern South America (to 
Brazil, Bolivia, and Peru). It is readily characterized by the ovate-oval leaves 
with bases prominently cuneately decurrent upon the petiole. 

AMAZONAS. Bagua, near Campamento Stte. Montenegro, 450 m, Wurdack 
1840 (F, GH, US). AYACUCHO. Aina between Haunta and Rio Apurimac, 
750-1,000 m, Killip & Smith 22793 (GH, US). CUZCO. Quispicanchis: Maria 
Patai, Cadena, 1,100 m, Vargas 11677 (US). HUANUCO. Huamalies, Monzon, 
Weberbauer 3442 (GH), Huanuco: downriver from Tingo Maria, 7,000 ft, Belshaw 
3104 (F, GH, MO, US), Fundo Honolulu, cerca a Tingo Maria, carretera 
Huanuco-Tingo Maria, 600-700 m, Ferreyra 2258 (MO, US, USM), Huanuco: 
Tingo Maria, shore of Rio Monzon, Asplund 12974 (F), Pozuzo, 800-900 m, 
Weberbauer 34376 (F, MO), Divisoria, cerca a Chinchono, entre Tingo Maria y 
Pucallpa, 1,500-1,600 m, Ferreyra 4314 (US) and 4292 (MO, US, USM). LORETO. 
Maynas: Florida, Rio Putumayo, at mouth of Rio Zubineta, 180 m, Klug 2172 (F, 
GH, MO), Rio Yahuasyacu, Rio Ampiyacu, Pebas, Revilla 713 (MO), Rio Am- 
piyacu, Pebas and vicinity, Plowman et al. 7268 (F), Dtto. Alto Nanay, Rio 
Nanay just below mouth of Quebrada Anguilla, McDaniel et al. 21503 (AMAZ, 
F, FSU, GA, IBE, MO, NATC, NY, RB, US, USM), Dtto. Iquitos, Quebrada de 
Aucaya, trocha de Nuevo Union, McDaniel & Rimachi 17171 (IBE, NATC), along 
Rio Amazonas S of Iquitos, Croat 19307 (GH, IBE), Iquitos, ca. 100 m, Killip & 
Smith 27386 (F, GH, US), Iquitos, San Juan, 120 m, Williams 3721 (F, GH), 
Iquitos, 120 m, Williams 8197 (F, GH), 5 m S of Iquitos, Croat 18144 (IBE), lower 
Rio Nanay, Williams 542 (F), Rio Nanay near Iquitos, between Bellavista & Sta. 
Clotilde, 110 m, McDaniel 13576 (IBE, MO), La Carretera de Momoncillo near 
Rio Momon, McDaniel & Rimachi 17007 (IBE); Ucayali: Contamana, near road to 
Oriente, 160-180 m, McDaniel 14057 (IBE). MADRE DE DfOS. Cocha Cashu, 
forest in vicinity of ox-bow lake of Rio Manii, between Panagua & Tayakome, 
Foster et al. 3488 (IBE). SAN MARTfN. Zepelacio, near Moyobamba, 1,200-1,600 
m, Klug 3412 (US), Mariscal Caceres: Tocache Nuevo, Quebrada Luis Salas (5 km 
NE Puerto Rizana), Schunke V. 6584 (IBE, MO), Mariscal Caceres: Tocache 
Nuevo, Fundo Jeroglifico, 400 m, Schunke V. 8282 (MO). 

28. Mikania haenkeana DC., Prodr. 5: 196. 1836. TYPE: "in montanis 
Peruviae," 1790, Haenke 142 (G-DC, holotype, not seen, IDC microfiche 800. 
820: 1.7; F, isotype). 

M. pellucidivenia Hieron., Bot. Jahrb. Syst. 36: 474. 1905. TYPE: Peru, near Tambillo, 
Jelski 600 & 642 (both B, destroyed). 

M. monzonensis Hieron., Bot. Jahrb. Syst. 40: 391. 1908. TYPE: Peru, Huamalies, Weber- 
bauer 3430 (B, holotype, destroyed, GH, photo; F, isotype). 

Herbaceous twiners; stems terete, glabrate, striate, internodes to 25 cm or more long. 
Leaves ovate, to ca. 12 cm long and 11 cm wide, margins entire, apices caudate- 
acuminate, bases cordate, upper surfaces glabrate, palmately 5-nerved, lower surfaces 
glandular-punctate, lighter than above; petioles to ca. 7 mm long, thin, puberulent. 
Heads sessile, disposed in somewhat elongate spiciform glomerules, these arranged in 
panicles; exterior bracts linear, 3-3.5 mm long, puberulent; involucral bracts lance- 
oblong, ca. 3 mm long, puberulent, apices acute; corollas ca. 3 mm long, greenish-white, 
tube scarcely distinguishable from and gradually expanding into the throat, teeth deltoid, 



^V'7*K/^P?3 ^jk.S . 

\\lm-lliji 
\ $i i * fip/ 

3^R irJ?:JW 




2 rnrn 



= 4 cm 



FIG. 3. Mi'Jtflnifl gwaco H. & A. A, habit; B, head; C, flower. (After McDaniel et al. 21503 
[IBE].) 



25 



26 FIELDIANA: BOTANY 

ca. 0.3mm long. Achenesca. 1.5mm long, black, somewhat hispid at the summit; pappus 
bristles ca. 30, white, 2.5-3 mm long, barbellate, thickened at the tips. 

Mikania haenkeana occurs at ca. 750-1,700 m elevation in Bolivia, Peru, 
Argentina (Salta), and Ecuador. It is characterized by the paniculate in- 
florescence with the heads sessile and disposed in somewhat spiciform elongate 
glomerules. 

HUANUCO. Pampayacu, hacienda at mouth of Rio Chinchao, 3,500 ft, Mac- 
bride 5064 (F, GH, US). JUNfN. La Merced, Hacienda Schunke, 4,000 ft, Mac- 
bride 5271 (F, GH, US), 5425 (F, GH, US), and 5810 (F, GH); La Merced, Soukup 
1537 (US). 

29. Mikania hexagonocaulis Holmes & McDaniel, Phytologia 471. 1977. TYPE: 
Peru, San Martin, Palo Blanco, al oeste del puente, Mariscal Caceres, Tocache 
Nuevo, December 16, 1972, Schunke V. 5734 (F, holotype). 

Climbing lianas; stems obscurely hexagonal to somewhat terete at first, later becoming 
distinctly hexagonal, reddish-brown, finely puberulent to somewhat scurry. Leaves 
ovate-elliptic, semifleshy, intense green when alive, to ca. 14 cm long and 6 cm wide, 
margins entire, apices attenuate, bases rounded, pinnately nerved, the nerves prominent, 
branching from the midvein and closely following same until separation, above finely 
puberulent, below lighter, glandular, somewhat puberulent; petioles 1.5-2 cm long. In- 
florescence a corymb, ca. 8 cm wide and 7 cm long; branchlets angular, densely puberu- 
lent. Heads sessile, temately arranged, ca. 12 mm long; exterior bracts subulate, ca. 1 mm 
long, glandular, weakly puberulent; involucral bracts greenish-yellow, the outer oblong, 
ca. 6 mm long, glandular, lightly puberulent, apices rounded with ciliate margins, inner 
narrowly obovate, ca. 7 mm long, glandular, puberulent, apices obtuse, margins ciliate; 
corollas narrow, white, ca. 5.5 mm long, throat scarcely distinguishable from the tube, 
teeth ca. 0.8 mm long, narrowly triangular. Achenes ca. 4.5 mm long, gradually enlarging 
from base to apex, olivaceous; pappus bristles 60-70, white, barbellate, ca. 6. 5 mm long. 

Mikania hexagonocaulis, known only from the type, is characterized by ter- 
nately disposed sessile heads, hexagonal stems, pinnately veined leaves, and 
subulate exterior bracts. 

30. Mikania hookeriana DC., Prodr. 5: 195. 1836. TYPE: Guyana, Schomburgk 
479 (G-DC, holotype, not seen, IDC microfiche 800. 820:1.3, F, photo; K, 
isotype, not seen). 

M. badieri DC, Prodr. 5: 194. 1836. TYPE: Guadeloupe, Badier (G-DC, holotype, not 

seen, IDC microfiche 800. 819:11.6). 
M. platyphylla DC., Prodr. 5: 195. 1836. TYPE: Peru, Poeppig2184 (G-DC, holotype, not 

seen, IDC microfiche 800. 819:111.7, F, photo). 
M. imrayana Griseb., Fl. Brit. W. Ind. 363. 1861. TYPE: Dominica, Imray (K, holotype, 

not seen). 

Willoughbya imrayana (Griseb.) Kuntze, Rev. Gen. PI. 1: 372. 1891. 
W. platyphylla (DC.) Kuntze, Rev. Gen. PI. 1: 372. 1891. 
Mikania badieri DC. var. kittsiana Urban, Symb. Antill. 5: 220. 1907. TYPE: St. Kitts, 

Belmont, Britton & Cowell 385 (US, holotype). 
M. vitrea B. L. Robinson, Contr. Gray Herb. 61: 22. 1920. TYPE: Venezuela, Colonia 

Tovar, Aragua, Fendler 2349 (GH, holotype; G, not seen, PH, isotypes). 
M. hookeriana DC. var. badieri (DC.) B. L. Robinson, Contr. Gray Herb. 104: 52. 1934. 
M. hookeriana DC. var. cardiophylla B. L. Robinson, Contr. Gray Herb. 104: 53. 1934. 

TYPE: St. Vincent, Smith 1090 (GH, holotype). 
M. hookeriana DC. var. kittsiana (Urban) B. L. Robinson, Contr. Gray Herb. 104: 52. 

1934. 
M. hookeriana DC. var. platyphylla (DC.) B. L. Robinson, Contr. Gray Herb. 104: 52. 

1934. 



MACBRIDE: FLORA OF PERU 27 

M. sanjacintensis Badillo, Bol. Soc. Venez. Ci. Nat. 10: 298. 1946. TYPE: Venezuela, 
Montana de San Jacinto, N side of Rio Chama, Merida, Steyermark 56665 (YEN, 
holotype, not seen; F, isotype). 

M. hookeriania DC. var. crassicaulis Steyermark, Fieldiana, Bot. 28: 658. 1953. TYPE: 
Venezuela, Ptarf-tepui, Bolivar, Steyermark 60015 (F, holotype). 

Herbaceous to woody lianas; stems terete, striate, often fistulose, glabrous to puberu- 
lent, intemodes 5-20 cm long. Leaves ovate to broadly ovate, to ca. 15 cm long and 10 cm 
wide, margins entire, often with prominent basal angles, apices acute to acuminate, bases 
rounded to slightly cuneate, upper surfaces glabrate to puberulous, often somewhat 
muricate, subpinnately 5-nerved from near the base, lower surfaces glabrate to puberu- 
lent; petioles to 4 cm long, thin. Inflorescence paniculate, ca. 12 cm in diameter and 12 cm 
high; heads disposed in rather dense short spikes or glomerules; branchlets terete to 
angular, puberulent. Heads sessile, 8-9 mm long; exterior bracts lance-linear, ca. l /2 the 
length of the involucre; involucral bracts oblong, ca. 5 mm long, apices rounded, puberu- 
lent, bases swollen; corollas 4-4.5 mm long, white, tube ca. 1.5 mm long, throat narrowly 
campanulate, 2-2.5 mm long, teeth ca. 0.5 mm long. Achenes 2.5-3.5 mm long, lightly 
glandular, brownish, ribs lighter in color; pappus bristles ca. 40, white, 5-6 mm long, 
thin, barbellate. 

Mikania hookeriana is a very widespread and highly variable species occurring 
from southern Mexico throughout Central America to Peru, Bolivia, and Brazil. 
An excellent account of the variation and synonymy of this species is presented 
by Robinson (1934). This species is distinguished by its sessile heads disposed 
in short spikes or somewhat elongate glomerules. The swollen bases of the 
involucral bracts are also a very useful diagnostic character. 

AMAZONAS. Lugar Aintami, monte al lado de Cenepa, Kayap 348 (MO), 
Quebrada Shimpunts, alto Rio Cenepa, monte al lado de Shimpunts, 800-1,100 
ft, Ancuash 29 (IBE), Kayap 426 (IBE, MO). HUANUCO. Pachitea: Honoria, 
Bosque Nacional de Iparia, Rio Pachitea, Schunke V. 1145 (US), Tingo Maria, km 
137-138, Carretera Huanuco-Pucallpa, 700 m, Ferreyra 906 (GH, MO, US, USM), 
Carretera a Pucallpa, km 210, Ridoutt s. n. (MO, USM), Agua Blanco (Carretera 
Monzon), Schunke V. 5868 (F). LORETO. Fortaleza near Yurimaguas, 140 m, 
Klug2897 (F, GH, US), Pampayacu, between Balsapuerto and Moyobamba, Klug 
3205 (US). SAN MARTfN. Zepelacio, near Moyobamba, 1,200-1,600 m, Klug 
3460 (F, GH, MO, US), Mariscal Caceres: Tocache Nuevo, camino a Shunte, 
Schunke V. 7344 (MO), Mariscal Caceres: Tocache Nuevo, aeropuerto de To- 
cache, 400 m, Schunke V. 3761 (F, US), Rioja, sabana, 1,000-1,100 m, Ferreyra 
18470 (MO, USM). 



31. Mikania huanucoensis Holmes & McDaniel, Phytologia 41: 190. 1979. 
TYPE: Peru, Huanuco, Rio Negro, pampa of fern, 1,000 m, January 14, 1961, 
Woytkowski 6192 (MO, holotype). 

Suffrutescent lianas to 10 m tall; stems striate-sulcate, fistulose, glabrous, internodes to 
4 cm long. Leaves elliptic, to ca. 5.5 cm long and 2.5 cm wide, margins entire-revolute, 
apices acute to acuminate, bases cuneate, upper surfaces very lightly sericeous toward the 
margins, obscurely pinnately nerved, with 2 pairs of secondary nerves arising within the 
basal half of the leaf, these arching toward the apex, lower surfaces glabrate to lightly 
pilose, the major nerves raised; petioles 0.5-0. 7 cm long, lightly sericeous. Inflorescence a 
panicle, to 14cm tall and 8cm wide, the ultimate branchlets spicate, opposite, to ca. 8cm 
long; branchlets striate, lightly glandular. Heads 10-11 mm long, loosely disposed; ex- 
terior bracts linear to narrowly elliptic, to ca. 1.5 mm long, glandular; involucral bracts 
oblong, ca. 6 mm long, glabrate, glandular, apices rounded, puberulent; corollas ca. 5 mm 
long, white, glandular, tube ca. 3 mm long, gradually expanding into the throat, throat ca. 
1.5 mm long, teeth obtuse, ca. 0.5 mm long, glandular. Achenes ca. 4.5 mm long, glandu- 



28 FIELDIANA: BOTANY 

lar toward the summit, dark olivaceous, ribs white; pappus bristles 40-45, ca. 5 mm long, 
barbellate, somewhat thickened toward the tips. 

Mikania huanucoensis, known only from the type, is characterized by its spi- 
cately disposed inflorescence with the heads 10-11 mm long. 

32. Mikania incasina B. L. Robinson, Contr. Gray Herb. 64: 11. 1922. TYPE: 
Peru, Amazonas, Chachapoyas, Mathews s. n. (K, holotype, not seen). 

Somewhat shrubby twiners; stems terete, striate, puberulent when young, soon gla- 
brate, intemodes to 14 cm long. Leaves triangular-ovate, to ca. 18 cm long and 9 cm wide, 
margins entire, apices acuminate, bases truncate to rounded, upper surfaces glabrous, 
3-5-nerved from the base, lower surfaces tomentose, the major nerves exserted from 
surface, prominent; petioles to 1.5 cm long, puberulent. Inflorescence a panicle, ca. 10 cm 
long and 10 cm wide; branchlets terete, puberulent; pedicels 3-5 mm long, puberulent. 
Heads ca. 10 mm long; exterior bracts linear, ca. 3 mm long, sparsely puberulent, gener- 
ally borne near the summit of the pedicel, but sometimes placed well beneath; involucral 
bracts oblong, ca. 6 mm long, somewhat glandular, apices obtuse, lightly puberulent; 
corollas 5 mm long, white, tube ca. 3 mm long, thin, throat campanulate-rotate, ca. 0.5 
mm long, teeth ovate, ca. 1.5 mm long, with 1 marginal pair of veins and another pair 
inward and parallel to the margin. Achenes ca. 4 mm long, olivaceous with white ribs; 
pappus bristles 55-60, white, ca. 5.5mm long, barbellate. 

Mikania incasina, known only from Peru, is characterized by its rotate- 
campanulate corolla throat with the teeth greater in length than the throat. 

HUANUCO. Carpish, entre Huanuco y Tingo Maria, 2,800-2,900 m, Ferreyra 
2084 (MO, US, USM). JUNfN. La Merced, Hacienda Schunke, ca. 4,000 ft, Mac- 
bride 5778 (F, GH, US). 

33. Mikania infesta B. L. Robinson, Contr. Gray Herb. 104: 37. 1934. TYPE: 
Peru, Junin, Pichis Trail, Yapas, alt. 1,350-1,600 m, June 28-29, 1929, Killip & 
Smith 25439 (US, holotype). 

Woody vines, stems terete, scabrid, internodes to 7 cm long. Leaves elliptic, to ca. 20 cm 
long and 7.5 cm wide, margins cuspidate-denticulate, apices acute-acuminate, bases ob- 
tuse to rounded, upper surfaces scabrous, pinnately veined, lower surfaces lightly pilose; 
petioles 3-3.5 cm long, somewhat scabrous. Inflorescence a panicle, ca. 8 cm in diameter, 
the ultimate segments spicately disposed; branchlets terete, scabrid. Heads ca. 5 mm 
long; exterior bracts lance-ovate, ca. 1.5 mm long, apices acute, somewhat ciliolate; in- 
volucral bracts oblong-ovate, 3.5-4 mm long, sparingly pilose, apices obtuse; corollas ca. 3 
mm long, greenish-white, tube ca. 1.5mm long, throat 1 mm long, teeth triangular, ca. 0.5 
mm long, pinkish. Achenes immature; pappus bristles ca. 25, white, ca. 4 mm long, 
scabrid, thickened at tips. 

Mikania infesta, known only from the type, is readily characterized by the 
spicately disposed heads and scabrid leaves. 

34. Mikania inornata B. L. Robinson, Contr. Gray Herb. 73: 26. 1924. TYPE: 
Peru, Ancash, Pampayacu, hacienda at mouth of Rio Chinchao, ca. 3,500 ft, 
July 19-25, 1923, Macbride 5061 (F, holotype). 

Herbaceous to slightly woody vines; stems somewhat hexagonal, glabrous, intemodes 
5-23 cm long. Leaves oblong-ovate, to ca. 10 cm long and 5.5 cm wide, margins entire, 
apices acuminate, bases acute to somewhat rounded, surfaces glabrous, 3(-5)-nerved 
from the base, reticulating veins obscure; petioles to ca. 4 cm long, glabrous. In- 
florescence a lax panicle to ca. 12 cm long and 6 cm in diameter; branchlets hexagonal, 
glabrous; pedicels 1-2 mm long, angular, glabrous. Heads ca. 8 mm long; exterior bracts 
oblong-ovate, 5-6 mm long, glabrate, apices acute, bases cuneate, margins somewhat 
ciliate, surfaces reticulate; involucral bracts ovate-oblong, 4.5-5.5 mm long, glabrate, 



MACBRIDE: FLORA OF PERU 29 

apices obtuse; corollas ca. 5 mm long, tube ca. 2.5 mm long, throat campanulate-rotate, ca. 
0. 7 mm long, teeth ca. 1.8 mm long, ovate, with 2 pairs of veins, 1 pair marginal, the other 
inward and paralleling the first. Achenes ca. 3 mm long; pappus bristles ca. 100, white, ca. 
5 mm long, barbellate, thickened at the tips. 

Mikania inornata, known only from the type, is similar in many respects to M. 
montana B. L. Robinson, but easily distinguished by its ovate corolla teeth with 
two pairs of veins. Mikania montana has linear corolla teeth with one pair of 
veins. 



35. Mikania iquitosensis B. L. Robinson, Contr. Gray Herb. 104: 38. 1934. 
TYPE: Peru, Loreto, Iquitos, woods, ca. 100 m, August 3-11, 1929, Killip & 
Smith 27080 (GH, holotype; F, isotype). 

Woody vines; stems terete, costate, with solid pith, glabrate, intemodes to 4 cm long. 
Leaves oblong, to ca. 8 cm long and 3.5 cm wide, margins entire, apices acute-acuminate, 
bases acute to obtuse, upper surfaces glabrate, pinnately nerved, lower surfaces glabrate; 
petioles to ca. 8 mm long, puberulent. Inflorescence a lax panicle; bracts linear, to ca. 1 cm 
long, puberulent; branchlets angular-costate, puberulent; pedicels ca. 2 mm long. Heads 
ca. 7 mm long; exterior bracts lance-linear, 1.5-2 mm long, puberulent; involucral bracts 
lance-linear, ca. 5 mm long, glabrate, apices acute-acuminate, slightly erase, bases 
slightly enlarged; corollas ca. 5 mm long, greenish-white, tube ca. 1.7 mm long, throat 
campanulate-turbinate, ca. 2.3 mm long, teeth deltoid, ca. 1 mm long. Achenes ca. 3 mm 
long, brownish; pappus bristles ca. 40, white, ca. 5 mm long, barbellate, distinctly thick- 
ened at the tips. 

Mikania iquitosensis is known only from the type. In many respects it is similar 
to M. decora Poepp., but lacks the distinctly racemosely disposed heads and 
prominently reticulate leaves of the latter species. The leaves of M. iquitosensis 
apparently turn dark brown to blackish upon drying. See M. shushunensis for 
distinctions between these two similar species. 



36. Mikania jelskii Hieron., Bot. Jahrb. Syst. 36: 475. 1905. TYPE: Peru, 
Cajamarca, near Cutervo, Jelski 634 (B, holotype, destroyed, F, photo). 

Somewhat woody lianas; stems terete, costate, puberulent, internodes to ca. 8cm long. 
Leaves elliptic, to ca. 7 cm long and 3 cm wide, margins entire, apices somewhat at- 
tenuate, bases rounded, upper surfaces glabrous, the major nerves arising from near the 
base, lower surfaces glabrous; petioles ca. 1.2 mm long, puberulent. Inflorescence 
paniculate, heads spicately disposed; branchlets puberulent; pedicels barely perceptible. 
Heads 5-7 mm long; exterior bracts ca. Va the length of the involucre, puberulent; in- 
volucral bracts oblong, 3-3.8 mm long, puberulent, inner surfaces pearly white, apices 
narrowed to an obtuse tip; corollas ca. 2.5 mm long, tube ca. 0.5 mm long, somewhat 
bulbous at the base, throat ca. 1.5 mm long, teeth deltoid, ca. 0.5 mm long. Achenes ca. 
1.5 mm long, black; pappus bristles 30-40, brownish, ca. 3 mm long, barbellate, thickened 
at the tips. 

Mikania jelskii is known only from Peru at 1,800-2,700 m elevations. It is very 
similar to M. oreopola, but the former can be distinguished by its dense in- 
florescence with slightly pedicellate heads, puberulent involucral bracts dark- 
ened on the outer surface and pearly white inside and pappus bristles with 
distinctly thickened tips. 

AMAZONAS. Bongora: Hills WNW of Pomacocha, 2,300-2,700 m, Wurdack 
929 (US); Chachapoyas, Mathews 1371 (GH, photo & frag.). CUZCO. Vill- 
cabamba, hacienda on Rio Chinchao, 6,000 ft, Macbride (F, GH, US). JUNlN. 
Huacapistana, 1,800 m, Killip & Smith 24292 (GH). 



30 FIELDIANA: BOTANY 

37. Mikania juninensis Holmes & McDaniel, Phytologia 42: 185. 1979. TYPE: 
Peru, Junin, Pichis Trail, Porvenir, 1,500-1,900 m, July 3-4, 1929, Killip & 
Smith 25912 (GH, holotype; F, isotype). 

Subligneous vines; stems terete, velvety tomentulose, internodes ca. 7 mm long. Leaves 
ovate, ca. 8.5 cm long and 4.5 cm wide, margins entire, apices acute to rounded, bases 
rounded to obtuse, upper surfaces glabrate, pinnately nerved, the major nerves with 
dense white appressed pubescence, lower surfaces crisped-hairy (mainly on the nerves), 
the veins exserted, lighter than above; petioles 1-2 cm long, velvety tomentulose. In- 
florescence a rather open panicle with heads clustered near the tips of the oppositely 
borne branchlets, ca. 30 cm long and 15 cm in diameter; branchlets terete, velvety tomen- 
tulose; pedicels 2-4 mm long, terete, velvety tomentulose. Heads 4-5 mm long; exterior 
bracts ovate-oblong, 1-1.2 mm long, glabrate, apices rounded, irregularly ciliate, borne 
well beneath the involucre; involucral bracts oblong, 3 mm long, glabrate, apices 
rounded; corollas 3-3.3 mm long, greenish-white, tube 1-1.2 mm long, gradually ex- 
panded into the turbinate throat, throat 1-1.2 mm long, teeth lanceolate to narrowly 
triangular, ca. 1.3 mm long. Achenes ca. 2 mm long, olivaceous; pappus bristles ca. 30, 
white, ca. 3 mm long, barbellate. 

Mikania juninensis is known only from two collections from Junin, the type 
and the one listed below. It is characterized by open panicles with heads some- 
what clustered toward the tips of the branchlets. 

JUNlN. Pichis Trail, Dos de Mayo, 1,700-1,900 m, Killip & Smith 25797 (GH). 

38. Mikania klugii B. L. Robinson, Contr. Gray Herb. 104: 38. 1934. TYPE: 
Peru, Loreto, Florida, Rio Putumayo, at mouth of Rio Zubineta, 180 m, 
May-July, 1931, King 2135 (GH, holotype; F, MO, isotypes). 

Subligneous lianas; stems terete, costate-striate, puberulent to crisped puberulent (es- 
pecially on younger parts), internodes ca. 5 cm long. Leaves ovate (only bracteal leaves 
available), to ca. 8 cm long and 5 cm wide, margins entire, apices long-attenuate, bases 
rounded to truncate to shortly cuneate, surfaces glabrate, prominently reticulate, sub- 
pinnately nerved from near the base; petioles to ca. 2 cm long, sulcate. Inflorescence a 
panicle, the ultimate branches spicately disposed, to ca. 30 cm long and 20 cm wide; 
branchlets crisped puberulent. Heads sessile, ca. 6 mm long; exterior bracts elliptic-ovate, 
0.5-1.0 mm long; involucral bracts elliptic-oblong, ca. 4 mm long, apices acute to a slightly 
rounded puberulent tip; corollas ca. 3.5 mm long, tube ca. 2 mm long, throat narrowly 
campanulate, ca. 1.0 mm long, teeth acute, ca. 0.5 mm long. Achenes (immature) ca. 1.8 
mm long; pappus bristles ca. 30, white, 4-4.5 mm long, barbellate, often thickened at the 
tips. 

Mikania klugii, known only from the type, is difficult to distinguish from M. 
aquaria. The exterior bract of M. aquaria is around one-half the length of the 
involucre, that of M. klugii one-fourth the length of the involucre. Klug 2960, 
cited with M. klugii in the original description has, among other differences, 
verticillate leaves and is referred to M. simpsonii. 

39. Mikania lanceolata Hieron., Bot. Jahrb. Syst. 29: 17. 1900. TYPE: Ecuador, 
"in sylvis temperatis prope Niebli." July 1873, Sodiro 7/1 (B. holotype, de- 
stroyed). 

Climbing shrubs; stems terete, glaucous-green to purplish, glabrate, internodes to ca. 9 
cm long. Leaves lanceolate, subcoriaceous, to ca. 12 cm long and 3 cm wide, margins 
entire, apices attenuate, bases rounded to cuneate, upper surfaces glabrate, reticulate, 
pinnately nerved, lower surfaces glabrate, lighter than above, prominently reticulate, 
veins exserted. Inflorescence a panicle; branchlets puberulent; heads arranged in 3's at 
the tips of the branches. Heads ca. 6mm long, sessile; exterior bracts subulate, ca. 0.5mm 



MACBRIDE: FLORA OF PERU 31 

long; involucral bracts spatulate, ca. 3 mm long, lightly puberulent, glandular, apices 
rounded; corollas ca. 4 mm long, glandular, tube ca. 1.5 mm long, the base slightly 
expanded, throat turbinate-subcylindric, ca. 2mm long, teeth triangular, ca. 0.5mm long. 
Achenes ca. 2 mm long, black, ribs lighter in color; pappus bristles ca. 30, white, ca. 4 mm 
long, barbellate. 

Mikania lanceolata is a very distinctive species of Peru and Ecuador rec- 
ognized by lanceolate leaves and sessile, ternately disposed heads ca. 6 mm 
long. 

CAJAMARCA. Tambillo, Jelski 601, 604, and 746 (all US). 

40. Mikania lancifolia B. L. Robinson, Contr. Gray Herb. 61: 17. 1920. TYPE: 
Peru, Puno, between Yuncacoya and Tambo Cachicachi on the way from 
Sandia to Chunchusmayo, 1,800-2,200 m, Weberbauer 1137 (B, holotype, de- 
stroyed, F, photo). 

Twining vines or somewhat shrubby; stems angular to terete, yellow-villous to gla- 
brate, internodes to 8 cm long. Leaves lanceolate, to ca. 10 cm long and 3.5 cm wide, 
margins remotely callous-denticulate, the teeth ca. 1 cm apart, apices attenuate, bases 
acute to obtuse, upper surfaces glabrate, rugulose, prominently reticulate, feather-veined, 
lower surfaces glandular, reticulate, veins exserted; petioles to ca. 1.5 cm long, thin. 
Inflorescence a lax panicle, to ca. 10 cm in diameter; branchlets pilose; pedicels 3-4 mm 
long, slender, lightly pilose. Heads ca. 7 mm long; exterior bracts lance-linear, ca. 2.5 mm 
long, borne at the summit of the pedicel; involucral bracts linear-oblong, ca. 4.5 mm long, 
glabrate except for the somewhat pilose acute apices; corollas ca. 3 mm long, white, tube 
ca. 1 mm long, throat campanulate, ca. 1 mm long, teeth triangular, equaling the throat in 
length. Achenes ca. 3 mm long, olivaceous; pappus bristles ca. 35, white, ca. 3.5 mm long, 
barbellate. 

An excellent diagnostic character of Mikania lancifolia, endemic to Puno, Peru, 
is its lanceolate leaves with callous-denticulate margins. 

PUNO. Sandia: 3 km above Santo Domingo, 1,950 m, Metcalf 30644 (MO, 
US). 

41. Mikania lasiopoda B. L. Robinson, Contr. Gray Herb. 73: 27. 1924. TYPE: 
Peru, Junin, La Merced, 2,000 ft, August 10-24, 1923, Macbride 5567 (F, 
holotype). 

Woody lianas; stems terete, costate, setose, internodes to ca. 20 cm long. Leaves nar- 
rowly ovate-oblong, to ca. 14 cm long and 6 cm wide, margins entire, apices attenuate, 
bases cordate on mature leaves, acute on bracteal leaves, upper surfaces glabrate, pin- 
nately nerved, lower surfaces slightly pilose, lighter than above, dark punctate; petioles to 
ca. 5 mm long, setose-pilose. Inflorescence a panicle, to ca. 15 cm long and 8 cm wide; 
branchlets angular-costate, pilose; pedicels to ca. 3 mm long, pilose. Heads 5-6 mm long; 
exterior bracts ovate, to ca. 1.5 mm long, pilose; involucral bracts oblong, 3-3.5 mm long, 
glabrous, apices rounded; corollas ca. 3.5 mm long, whitish, tube ca. 1.4 mm long, throat 
campanulate, ca. 1.3 mm long, teeth triangular, ca. 0.8 mm long. Achenes ca. 2.5 mm 
long, glabrous, light green; pappus bristles 30-35, white, ca. 3.5 mm long, barbellate. 

Mikania lasiopoda, endemic to Junin, is marginally distinct from the highly 
variable M. banisteriae DC., but here maintained as a separate species with 
much smaller heads (5-6 mm long), involucral bracts of 3-3.5 mm long, and 
smaller corollas and achenes. Older stems lack the dense pubescence normally 
present on M. banisteriae. 

JUNlN. La Merced, Hacienda Schunke, 400 ft, Macbride 5729 (F, GH, US). 



32 FIELDIANA: BOTANY 

42. Mikania leiostachya Benth., PI. Hartw. 201. 1845. TYPE: Colombia, Cun- 
dinamarca, in descent from Guaduas to Honda, Hartweg 1120 (K, holotype, 
not seen). 

Lianas; stems terete, glabrate to minutely puberulent. Leaves ovate, subcoriaceous, to 
ca. 24 cm long and 13 cm wide, margins entire, often slightly recurved, apices acuminate, 
bases rounded, surfaces glabrate, subpinnately 5-nerved from near the base, prominently 
reticulate, veins exserted; petioles to ca. 3.5 cm long. Inflorescence a panicle, the ultimate 
branches spicate; branchlets angled, puberulent. Heads sessile, 5-6 mm long; exterior 
bracts lance-ovate, ca. 1.5 mm long, apices acute-acuminate; involucral bracts ovate- 
oblong, 3-3.5 mm long, puberulent, apices obtuse to rounded; corollas 2.5-3 mm long, 
white, tube 1-1.5 mm long, throat 1-1.5 mm long, teeth triangular, ca. 0.5 mm long. 
Achenes ca. 1.5-2 mm long, dark; pappus bristles ca. 35, white, 3.5-4 mm long, barbel- 
late, thickened at the tips. 

A widely distributed species of Central America and northern South America 
characterized by subcoriaceous ovate leaves and spicately disposed heads. 

HUANUCO. Huanuco: Caymba, entre Huanuco y Tingo Maria, 150-250 m, 
ferreyra 4206 (MO, US, USM). DPTO. UNKNOWN. Chota, Huambos, ca. 2,000 
m, Soukup 4521 (US). 

43. Mikania lindleyana DC., Prodr. 5: 195. 1836. TYPE: "Guiana" (G-DC, 
holotype, not seen, IDC microfiche 800. 1820: I.I). 

Herbaceous vines; young stems angular, puberulent, soon glabrate, multicostate, 
internodes to 17 cm long. Leaves ovate, to ca. 15 cm long and 8 cm wide, margins entire, 
apices acuminate, bases rounded, surfaces glabrate, 5-7-nerved from the base; petioles to 
6 cm long, glabrate. Inflorescence a corymb, branching trichotomously, ca. 5 cm long and 
6 cm wide; branchlets angular, puberulent; pedicels to 6-7 mm long, angular, puberulent. 
Heads ca. 8 mm long; exterior bracts ovate, ca. 4 mm long, lightly puberulent to glabrate, 
apices acute, prominently nerved; involucral bracts ovate-oblong, ca. 5 mm long, lightly 
puberulent, apices rounded; corollas 3-4 mm long, tube and throat about the same length, 
throat campanulate, teeth triangular, slightly greater in length than the throat. Achenes 
ca. 3 mm long, olivaceous; pappus bristles ca. 4 mm long, carneous, ca. 60, barbellate, 
slightly thickened toward the tips. 

Mikania lindleyana of Brazil, the Guianas, Venezuela, Trinidad, and Peru, is 
characterized by the normally ternately branching corymb. 

AMAZON AS. Quebrada chichejam ensa, Rio Cenepa, 920 ft, Ancuash 583 
(IBE, MO). JUNfN. Pichis Trail, Yapas, 1,350-1,600 m, Killip & Smith 25482 (GH, 
US). LORETO. Iquitos, 100 m, Killip & Smith 27104 (GH, US); Balsapuerto, 220 
m, King 3060 (GH, MO, US). 

44. Mikania lutescens B. L. Robinson, Contr. Gray Herb. 80: 37. 1928. TYPE: 
Peru, Ayacucho, Huanta, Coimacola Valley, 2,900 m, February 28-March 10, 
1926, Weberbauer 7574 (F, holotype). 

Twining shrubs; stems subterete, glabrate, internodes 2-9 cm long. Leaves triangular, 
ca. 5 cm long and 3 cm wide, margins crenate-dentate to coarsely serrate to somewhat 
irregularly palmately lobed at the base, the teeth mucronate, apices acute-acuminate, 
bases cordate, surfaces glabrate, palmately 5-7-nerved; petioles ca. 2 cm long, thin, con- 
nected by a thin lacerate stipule-like ridge. Inflorescence a corymb, ca. 6 cm long and 6 cm 
wide; branchlets angular, somewhat winged, glabrous; pedicels to ca. 5 mm long, angu- 
lar, winged. Heads ca. 10 mm long; exterior bracts ovate, somewhat petiolate, to ca. 6 mm 
long, lightly pilose, apices acute; involucral bracts linear-lanceolate, 7-8 mm long, gla- 
brous, apices acute-acuminate, the center of each base expanded into a small knoblike 
protuberance; corollas ca. 5.2 mm long, reported as yellowish (see below), tube ca. 2.7mm 



MACBRIDE: FLORA OF PERU 33 

long, throat subcylindrical-campanulate, ca. 1 mm long, teeth narrowly triangular, ca. 1.5 
mm long. Achenes ca. 3 mm long, glabrous, brownish; pappus bristles ca. 60, white, ca. 7 
mm long, barbellate. 

Mikania lutescens, known only from the type, is in many respects similar to M. 
brachyphylla Hieron., which has somewhat coriaceous leaves usually less than 1 
cm long. Mikania lutescens has leaves ca. 5 cm long, membraneous in texture. 
Apparently the specific epithet refers to the pale yellow flowers. 

45. Mikania marinii Cabrera, Bol. Soc. Arg. Bot. 5: 37. 1953. TYPE: Peru, 
Cuzco, Calca, Manto, Lares, en borde de bosque a 2,400 m, August 1950, 
Marin 2360 (LP, holotype, not seen). 

Scandent vines; stems terete, multistriate, hazel-hispid, 2-3 mm thick, internodes 5-12 
cm long. Leaves ovate-lanceolate, 7-8 mm long and 2. 5-3. 5 cm wide, membranous, apices 
attenuate, bases rounded, margins remotely denticulate, teeth 0.1-0.3 mm long, pinnately 
5-nerved, minutely bullate, strigose-hispid, below brownish-pubescent; petioles 2-3 cm 
long, densely hazel-subretrorse-hispid. Inflorescence thyrsoid-paniculate, to ca. 15 cm 
long and 8 cm wide; pedicels 2-6 mm long, hispid. Heads large; external bracts widely 
lanceolate, hazel-hispid, 2-4 mm long; involucral bracts lanceolate, ca. 8 mm long and 2 
mm wide, apices attenuate, submucronate, outer involucral bracts somewhat hirsute, 
inner glabrous; corollas 8 mm long, teeth ca. 1 mm long. Achenes 4.5 mm long, glabrate to 
lightly sericeo-pubescent at the base; pappus bristles numerous, carneous, ca. 7 mm long. 

The above description is compiled from the original publication of Cabrera 
(1953), which includes an excellent illustration. This species, known only from 
the type, is characterized by hazel-colored pubescence on the young stems, 
leaves, and involucral bracts. 

46. Mikania mathewsii B. L. Robinson, Contr. Gray Herb. 61: 18. 1920. TYPE: 
Peru, without exact location, in the Andes, Mathews 1368 (GH, holotype, F, 
photo & fragm.; NY, isotype). Figure 4. 

M. tenax Sch.-Bip. ex B. L. Robinson, Contr. Gray Herb. 80:41. 1928. TYPE: Peru, in 
mountains near San Govan, July 1854, W. Lechler 2355 (K, holotype, not seen, GH, 
photo & fragm., not seen; G, not seen, BR, isotypes). 

Somewhat woody lianas; stems terete, puberulent, soon glabrate, the nodes generally 
puberulent, internodes to 6 cm long. Leaves ovate to ovate-oblong (bracteal leaves 
lance-ovate), to ca. 14 cm long and 9 cm wide, margins entire, apices acuminate, bases 
acuminate, upper surfaces glabrate, pinnately nerved, with 3 pairs of nerves separating 
within the basal Va of the leaf, lower surfaces paler than above, often punctate, glabrate, 
the nerves often puberulent; petioles to ca. 2 cm long, puberulent. Inflorescence a panicle 
with heads disposed in somewhat dense corymbiform clusters at the tips of the branches; 
branchlets appressed pubescent; pedicels to ca. 3 mm long, thin puberulent. Heads 6-7 
mm long; exterior bracts lanceolate to ovate, 1.5-2 mm long, margins ciliate, borne at the 
base of the pedicel; involucral bracts ovate-oblong, ca. 3.3 mm long, puberulent, espe- 
cially at the apex, narrowing to a somewhat tumid base, apices obtuse; corollas ca. 4 mm 
long, tube ca. 1.3 mm long, throat subcylindric-semicampanulate, ca. 2.1 mm long, teeth 
narrowly triangular, ca. 0.6 mm long. Achenes ca. 2.8 mm long, dark brown; pappus 
bristles ca. 30, white, ca. 4.5 mm long, barbellate. 

Mikania mathewsii is endemic to Peru at elevations up to ca. 1,200 m. It is 
characterized by a paniculate inflorescence with heads rather densely clustered 
near the tips of the branches. 

CUZCO. Quispicanchis: entre Oraya y 15 Mil, 730, Vargas 14597 (US), Quis- 
picanchis: Quincemil, 800 m, Marin 1558 (US), Pamaostamso, 970 m, Vargas 
11013 (US). HUANUCO. Tingo Maria, Rio Monzon, Ferreyra 10040 (MO, US, 



34 



FIELDIANA: BOTANY 




= 2 mm 



= 4 cm 



FIG. 4. Mikania matheivsii B. L. Robinson. A, habit; B, head; C, flower; D, leaf. (After 
McDaniel & Rimachi 21971 [IBE].) 

USM). LORETO. Between Balsapuerto and Moyobamba, 600-1,200 m, King 3205 
(F, GH, MO), Coronel Portillo: Padre Abad, granja del Sr. Barrera, 295 m, 
Schunke V. s. n. (F), Yurimaguas, lower Rio Huallaga, 135m, Killip & Smith 27556 
(F, GH, US) & 27949 (F, GH, US), Iquitos, 200 m, Killip & Smith 26968 (GH, US), 
Iquitos, Quisto Cocha, McDaniel & Persinos 10696 (IBE, NATC). SAN MARTfN. 
Mariscal Caceres: Tocache Nuevo, camino a Shunte, 500-800 m, Schunke V. 7342 
(MO), Rioja, sabana, 1,000-1,100 m, Ferreyra 18462 (MO, USM). 



47. Mikania mazanensis Holmes & McDaniel, Phytologia 31: 275. 1975. TYPE: 
Peru, Loreto, Maynas, Gamitanacocha, Rio Mazan, 100-125 m, February 12, 
1935, Schunke 231 (F, holotype; GH, UC, isotypes). 

Twining vines; stems obscurely angled, glabrous, internodes 6-14 cm long. Leaves 
ovate, 5-10 cm long, 3-6 cm wide, semicoriaceous, margins subentire to weakly crenate- 
undulate, apices acute to acuminate, bases subcordate to truncate, upper surfaces gla- 



MACBRIDE: FLORA OF PERU 



35 



brous, weakly muricate, pale green, palmately 5-nerved from the base, major nerves 
whitish, lower surfaces glabrous, weakly bullate, pale green, major nerves wide, to 2.5 
mm, whitish; petioles 2-3 cm long, glabrous to weakly puberulent, whitish, ca. 2 mm 
thick. Inflorescence corymbose, 3-8 cm long, 5-10 cm wide; branchlets angled, glabrous; 
pedicels 5-15 mm long, glabrous. Heads 7-10 mm long; exterior bracts subulate to linear, 
2-3 mm long, glabrous, borne near the middle or base of pedicel; involucral bracts lance- 
elliptic to linear lanceolate, ca. 7 mm long, glabrous to weakly puberulent; corollas sal- 
verform, 4.5-4.7 mm long, cream-colored, glabrous, tube ca. 2.5 mm long, glandular near 
throat, throat ca. 0.7 mm long, teeth 1.3-1.5 mm long, lanceolate, distinctly longer than 
the throat. Achenes (immature) ca. 2.5 mm long, smooth, light olive in color; pappus 
bristles ca. 120, in 2 series, reddish in age, ca. 5 mm long, remotely barbellate at base, 
distinctly barbellate at tips. 

Mikania mazanensis, known only from the type, is readily identified by its 
corymbose inflorescence with pedicels 5-15 mm long, the linear to subulate 
exterior bract borne well beneath the involucre, the mostly linear-lanceolate 
involucral bracts with involute margins, the corolla teeth distinctly longer than 
the throat and numerous pappus bristles in two series. 

48. Mikania megalophylla Holmes & McDaniel, Phytologia 31: 276. 1975. 
TYPE: Peru, San Martin, Rio Huallaga, S of Shapaja 1-4 km, 900 ft, July 28-30, 
1937, Belshaw 3153 (F, holotype; MO, isotype). Figure 5. 

Twining lianas to ca. 5 m long; stems terete, striate to sulcate, hollow, glabrous to 
villous (at node), intemodes to 10 cm or more. Mature cauline leaves elliptic, 33-40 cm 
long, 13-17 cm wide, margins entire to weakly and obscurely crenate, apices cuspidate to 
caudate-cuspidate, bases truncate to obtuse, upper surfaces glabrous, dull green, veins 



8 cm 





4 nun 



2 mm 



4 cm 



FIG. 5. Mikania megalophylla Holmes & McDaniel. A, inflorescence; B, head; C, flower; 
D, leaf. (A & D after Belshaw 3153 [F]; B & C after Foster 2467 [IBE].) 



36 FIELDIANA: BOTANY 

pinnate, exserted, prominent, lower surfaces glabrous, lighter than above, veins exserted, 
prominent, slightly puberulent on the major veins; petioles to ca. 4 cm long, glabrous, 
sulcate, thick. Inflorescence paniculate, ca. 20 cm long, 7-15 cm wide, open, ultimate 
branchlets spicate to glomerate-spicate, 1-4 cm long; branchlets slightly angular, puberu- 
lent. Heads sessile, 3.5-5 mm long, somewhat antrorse and paralleling branchlets (rather 
than being perpendicular to branchlets); exterior bracts elliptic to narrowly ovate, 2-4 mm 
long, glabrous to weakly pilose, margins entire to erase, apices acute to subobtuse; 
involucral bracts ovate, 3.5-4 mm long, pilose, margins entire, slightly involute, apices 
obtuse, with a tuft of hair; corollas salverform, 2.5-3 mm long, white, sparingly glandular, 
tube 1.25-1.5 mm long, throat ca. 1 mm long, teeth ca. 0.5 mm long, triangular. Achenes 
ca. 1.5 mm long, glandular, smooth, light to dark brown, with a darkened ring at the 
summit near pappus bristles; pappus bristles 30-35, white, ca. 3 mm long, barbellate, 
thickened at the tips. 

Mikania megalophylla grows at elevations to ca. 900 m in Acre, Brazil; 
Huanuco, Madre de Dios; and San Martin, Peru. Mature cauline leaves of this 
species are up to 40 cm long, the largest known for the genus. 

HUANUCO. Pendencia, in forest, 900 m, Woytkowski 7521 (MO). MADRE DE 
DlOS: Manu, forest behind settlement, Foster 2467 (IBE, F, USM). SAN MAR- 
TIN. Juan Jui, alto Rio Huallaga, 400-800 m, King 4363 (F, MO), W side of Rio 
Huallaga, S of Shapaja 1-4 km, 900 ft, Belshaw 3154 (F, MO, UC), Lamas, arriba 
de San Jose de Sisa, selva, 400-450 m, ferreyra 7914 (MO, USM). 

49. Mikania micrantha HBK. Nov. Gen. Sp. PI. 4: 105. ed. fol. 1818. TYPE: 
Venezuela, "prope Caripe Cumanesianum," Humboldt & Bonpland (P, 
holotype, not seen). Figure 6. 

Kleinia alata G. F. W. Mey., Prim. Fl. Esseq. 249. 1818. nomen confusum (apparently 

based upon more than one element). TYPE: not seen, probably nonexistent. 
Mikania orinocensis HBK., Nov. Gen. Sp. PI. 4; 105. ed. fol. 1818. TYPE: Venezuela, 

"insulae Pararuma, Missiones del Orinoco," Humboldt & Bonpland (P, holotype). 
M. alata (Mey.) DC., Prodr. 5: 197. 1836. (See Robinson, Contr. Gray Herb. 104: 57. 

1934.) 
M. subcrenata H. & A., Campanion Hot. Mag. 1: 243. 1836. TYPE: Argentina, woods of 

Tucuman, Tweedie 1189 (K, holotype, not seen). 
M. umbellifera Gardner, London]. Bot. 4: 109. 1845. TYPE: Brazil, Organs Mts., ca. 4,000 

ft, Gardner 483 (K, holotype, not seen). 
M. subcymosa Gardner, London J. Bot. 6: 448. 1847. TYPE: Brazil, Goias, Gardner 3271 

(K, holotype; NY, P, US, isotypes). 

M. scandens var. umbellifera (Gardner) Baker in Mart., Fl. Bras. 6: 249. 1876. 
M. glechomaefolia Schultz-Bip. ex Baker in Mart., Fl. Bras. 6: 249. 1876. TYPE: not 

specified. 
M. scandens var. hirsuta Hieron., Bot. Jahrb. Syst. 19: 47. 1894. TYPE: Colombia, Cauca 

prope Cali et Quilichao, Lehman 4888 (B, holotype, destroyed). 
Willoughbya micrantha (HBK.) Rusby, Mem. Torrey Bot. Club 4: 211. 1895. 
Mikania scandens var. villosa Hieron., Bot. Jahrb. Syst. 36: 473. 1905. SYNTYPES: Peru, 

Tambillo, jelski 600, 640, 641, & 695 (all B, destroyed). 

M. micrantha f. hirsuta (Hieron.) B. L. Robinson, Contr. Gray Herb. 64: 43. 1922. 
M. sinuata Rusby, Bull. N.Y. Bot. Card. 8: 127. 1912. TYPE: Bolivia, San Juan, March 20, 

1902, Williams 207 (NY, holotype, US, isotype). 

Herbaceous to semiwoody twiners; stems terete to angled, mostly glabrate, intemodes 
5-20 cm long. Leaves ovate, thin, 2-13 cm long and 3-10 cm wide, margins entire to 
crenate, undulate, or dentate, apices acute to acuminate, bases cordate, sinuses open, 
upper surfaces glabrous, 3-5(7)-nerved from the base, lower surfaces glabrous; petioles 
1-8 cm long, glabrous to pilose, slender. Inflorescence a rather large corymbose panicle, 
3-6 cm long, 3-10 cm wide; branchlets lightly winged, glabrous. Heads 4-6 mm long; 
exterior bracts lanceolate, 2-3 mm long, glabrous, apices acute; involucral bracts ovate- 
oblong to rarely elliptic, 3-4 mm long, glabrous, apices acute; corollas campanulate, 2.5-3 



MACBRIDE: FLORA OF PERU 



37 



mm long, white to greenish, lightly glandular, tube 1-1.5 mm long, throat ca. 1 mm long, 
teeth deltoid, ca. 0.5 mm long, recurved. Achenes 1.5-2 mm long, sparingly glandular, 
black; pappus bristles 33-36, white, 2-3 mm long, barbellate. 

Mikania micrantha is very widespread in disturbed areas, wet places, river- 
banks, etc., to ca. 3,000 m elevation; flowering all year. The species occurs in 
wet tropical North America (excluding Florida), the West Indies, and South 
America to northern Argentina and is widely naturalized in Southeast Asia, the 
East Indies, Sri Lanka, India, Hong Kong, Solomon Island, Guam, Fiji, etc. 

Because of the widespread natural distribution and presence of several highly 
variable and often confusing characters, Mikania micrantha is often confused 
with several other species of Mikania. These include M. scandens, which is 
largely confined to the United States, and M. congesta which has the in- 
florescence a dense subspherical glomerule. The best diagnostic characters for 
M. micrantha include the ovate leaves with cordate bases, large corymbose 
panicle, heads 4-6 mm long, involucral bracts normally less than 4 mm long, 
and whitish to greenish corollas with recurved teeth. 



2 nun 




2 nun 



FIG. 6. Mikania micrantha HBK. A, habit; B, head; C, flower. (After McDaniel 16221 
[IBE].) 



38 FIELDIANA: BOTANY 

AMAZONAS. Huampani, 600-800 ft, Berlin 1977 (IBE), Quebrada Huampani, 
600 ft, Kayap 1167 (MO), Rio Cenepa, La Banda, Huampani, 840 ft, Berlin 287 
(MO), Lugar Kayamas, monte orilla de Cenepa, Kayap 274 (GH, MO), Kusa, Rio 
Numpakin, 1,100-1,300 ft, Ancuash 60 (MO). ANCASH. Caracha (way from 
Supe to Ocros), Weberbauer 2638 (GH). AYACUCHO. Aina, between Haunta 
and Rio Apurimac, 750-1,000 m, Killip & Smith 23113 (F, GH, NY, US), Rio 
Apurimac valley, near Kimpitiriki, 400 m, Killip & Smith 22851 (GH, NY, US). 
CAJAMARCA. Tambillo, Jelski 640 (GH) and 642 (GH, US). CUZCO. Entre 
Salvacion y Rio Yunsayo, 620 m, Vargas 16286 (US), entre Sta. Rita y Machupic- 
chu, 2,000-2,100 m, Vargas 3356 (F), Callanga, en la purma, 1,500 m, Woytkowski 
491 (MO, USM). HUANUCO. Huanuco, 3,000 ft, Macbride 3227 (F, GH), 
Huanuco, ca. 7,000 ft, Macbride 3203 (F, GH), Pachitea: Honoria, en bosque alto 
en el camino a Ayamiria a 4 km, 300-400 m, Schunke V. 1528 (US), Pachitea: 
Honoria, Bosque Nacional de Iparia, Rio Pachitea, cerca 3 km del campamento 
Miel de Abeja, 300-400 m, Schunke V. 1836 (F, US), Moena, 600 m, Woytkowski s. 
n. (MO, USM). JUNlN. La Merced, 700 ft, Macbride 3203 (US) and 3227 (US). 
LIBERTAD. Cicama Valley, on road to Chiclin, Smith 71 (US), Trujillo, near sea 
level, Killip & Smith 21502 (F, GH, NY, US). LIMA. Barranca Hda. Molina, 320 
m, Vargas 4617 (F), Ridoutt s. n. (MO, USM), Chosica, 3,000 ft, Macbride (F, GH), 
Chosica, Zuriiga s. n. (MO, USM), Esposto s. n. (IBE, MO, USM), Surco, 2,000 m, 
Asplund 11035 (US), Lima, Gaudichaux s. n. (GH), Lima, Wilkes89 (GH, NY, US), 
near Lima, Nation 203 (K), Lima, Anderson s. n. (NY), Hacienda Patao, Barranca, 
camino a orillas del Rio Pativilca, Ridoutt s. n. (MO, USM), Nafta, a 20 km E de 
Lima, Ferreyra 10548 (MO, USM). LORETO. Balsapuerto, 220 m, Klug 2977 (F, 
GH, MICH, MO, NY, US), Maynas: Fernando Lores, Dos Unidos, Rio 
Amazonas below Capironal, big bend above Rio Tahuayo, McDaniel & Rimachi 
(F, IBE, MO), Maynas: Mazan, Baradero de Mazan, near Mazan, McDaniel & 
Rimachi (IBE, NATC), Maynas: Alto Nanay, Trocha a Santa Rosa, a 3 km de 
Santa Maria de Nanay, 130 m, Schunke V. 2502 (US), Alto Nanay, Rio Nanay just 
below mouth of Quebrada Anguilla, McDaniel et al. 21507 (AMAZ, F, IBE, MO, 
NATC, NY, RB, US, USM), Manfinfa on the upper Rio Nanay, Williams 1105 (F, 
GH), Maynas: Iquitos, Isla de Escabino near Santa Maria de Nanay, McDaniel & 
Rimachi (IBE, NATC), Maynas: Iquitos, Versailles, McDaniel 11744 (FSU, IBE, 
MO), lower Rio Nanay, Williams 492 & 523 (both F, GH), Rio Nanay just below 
Moronacocha near Iquitos, 120 m, Gentry & Daly 19125 (MO), lower Rio Hual- 
laga, 155-210 m, Williams 4703 (F, GH), Pebas on the Amazon River, Williams 
1860 (F, GH), La Victoria on the Amazon River, Williams 3009 (F, GH), Rio 
Maranon valley, 150 m, barranca between mouth of Rio Monora and Rio 
Pastaza, Killip et al. 29232 (NY, US), IVTTA, km 59, Pucallpa-Tingo Maria road, 
Gentry et al. 18591 (MO), Coronel Portillo: Cerca a Neshuya, IVITA, selva tropi- 
cal, 250 m, Ferreyra 17183 & 27563 (both MO, USM), Bosque Nacional von 
Humboldt, km 86, Pucallpa-Tingo Maria road, 300 m, Gentry et al. 18647 (MO), 
San Jorge, cerca Pucallpa, 280-300 m, Ferreyra 13006 (MO, USM). MADRE DE 
DfOS. Manu: Parque Nacional de Manii, several miles upriver from Cocha 
Cashii, Foster 2706 (IBE), Cocha Cashii, Foster et al. 3310 (IBE). PIURA: Huan- 
cabamba, entre Serran y Canchaque, 500-600 m, Ferreyra 10779 (MO, USM). 
SAN MARTIN: Juan Jui, alto Rio Huallaga, 400-800 m, Klug 4307 (F, GH, MO, 
NY, UC, US), Tarapoto-Yurimaguas Highway, km 14-17, 820-850 m, McDaniel 
13797 (IBE, NATC), Mariscal Caceres: Tocache Nuevo, Quebrada de Ishichimi, 



MACBRIDE: FLORA OF PERU 39 

Schunke V. 3912 (IBE), Moyobamba, Mathews 1836 (BR, K), San Roque, 1,350- 
1,500 m, Williams 7058 & 7746 (both F, GH), Zepelacio, near Moyobamba, 
1,200-1,600 m, Klug 3453 (F, GH, MO, NY, US). 

50. Mikania microptera DC., Prodr. 5: 196. 1836. TYPE: Brasil, Bahia, Blanchet 
1710 (G-DC, holotype, not seen, IDC microfiche 800. 820:11.2; MO, isotype). 

M. scandens (L.) Willd. var. microptera (DC.) Baker in Mart., Fl. Bras. 6: 259. 1876. 
M. hexagona B. L. Robinson, Proc. Am. Acad. 47: 196. 1911. TYPE: Venezuela, near 
Tovar, 1,700 m, 1854-55, Fendler626 (GH, holotype). 

Vigorous herbaceous twiners; stems hexagonal, winged, younger parts puberulent to 
pilose, soon glabrate, internodes 10-20 cm long. Leaves ovate to triangular, 5-15 cm long 
and 3-18 cm wide, margins coarsely serrate to dentate to crenate, often with two di- 
vergent, pointed lateral lobes, apices acute to attenuate, bases cordate, sinuses wide, 
open, broadly cuneate on petiole at point of insertion, upper surfaces pale green, gla- 
brous, 3-5-nerved from the base, lower surfaces lighter than above, glabrous, sometimes 
glandular, veins prominent, lightly pilose; petioles 3-8 cm long, weakly winged, glabrous 
to slightly villous. Inflorescence corymbose, terminal, 4-8 cm long and 5-12 cm wide; 
branchlets pilose, winged; pedicels 3-10 mm long, angled, winged, pilose. Heads 6-8 mm 
long; exterior bracts linear to lanceolate, ca. 3 mm long, pilose, apices acute to attenuate; 
involucral bracts narrowly elliptic to oblong, 5-6 mm long, lightly pilose, weakly nerved, 
margins thin, apices acute to rounded, often with mucronate tips; corollas ca. 4 mm long, 
white to greenish, sparingly glandular, tube ca. 2 mm long, throat ca. 0.5 mm long, teeth 
longer than the throat, lanceolate, ca. 1.5 mm long. Achenes 2.7-3.5 mm long, glandular, 
tan to brown; pappus bristles 40-50, white, ca. 3.5 mm long, barbellate, gradually thin- 
ning toward the tips. 

Mikania microptera occurs on riverbanks and in other wet places to ca. 1,000 
m; flowering July-February, occasionally at other times. It is known from wet 
tropical west and central Africa, and South America, the Guianas and western 
Venezuela, the Amazon Basin (Peru, Bolivia), and Amapa, Bahia, and Parana 
Brasil. Mikania microptera apparently is native to both continents, though it 
appears with more regularity in African collections, possibly indicating more 
extensive and thorough collecting in Africa than South America. This species is 
very readily identified by its distinctly hexagonal usually glabrate stems with 
narrow wings. 

LORETO. Maynas: Gamitanacocha, Rio Mazan, 100-125 m, Schunke 386 (F, 
US); Dtto. Iquitos: Rio Itaya, trocha de Sanangal, disturbed forest of restinga, 
McDaniel & Rimachi 23282 (AMAZ, DUKE, F, FLAS, FSU, GA, IBE, MO, NY, 
RSA, SMU, US, USM, VDB); Carretera de Zungaro Cocha, near Quebrada de 
Shushuna, McDaniel & Rimachi 23301 (ALA, AMAZ, DUKE, F, FLAS, FSU, GA, 
IBE, LACM, MO, NATC, NO, RB, SMU, TENN, US, USM, VDB); exact location 
unknown, Poeppig 2335 (P). SAN MARTfN. Tocache Nuevo, Quebrada de 
Huaquisha, margen izquierda del Rio Huallaga, 400 m, Schunke V. 7017 (MO). 

51. Mikania montana B. L. Robinson, Contr. Gray Herb. 73: 29. 1924. TYPE: 
Peru, Huacachi near Muna, 1,982 m, May 20-June 1, 1923, Macbride 4169 (F, 
holotype). 

Lianas; stems terete, somewhat succulent, glabrate, internodes 3-7 cm long. Leaves 
ovate, fleshy, to ca. 5 cm long and 3 cm wide, margins entire to somewhat cuspidate- 
denticulate, apices acute, bases rounded to an acute insertion, surfaces glabrous, 5-7- 
nerved from the base, reticulate-areolate; petioles to ca. 2 cm long, glabrate, connected by 
a stipule-like annular ring. Inflorescence a corymb, to ca. 8 cm in diameter; branchlets 
angled, puberulent; pedicels to ca. 4 mm long, puberulent. Heads ca. 10 mm long; exterior 



40 FIELDIANA: BOTANY 

bracts ovate-lanceolate, ca. 1 mm long, prominently reticulate, apices acute, borne at the 
top of the pedicel; involucral bracts oblong, ca. 8 mm long, glabrate except for the ciliated 
acute apices; corollas 5.5 mm long, greenish to yellow-green, tube ca. 3 mm long, throat 
ca. 0.5mm long, teeth linear, ca. 2.5mm long. Achenesca. 4.5mm long, glabrous; pappus 
bristles ca. 80, white, ca. 5 mm long, barbellate. 

Mikania montana is thus far known only from Peru at elevations above 1,900 
m. 

CUZCO. La Convencion, on steep dry ridges ca. Vz-way between Camp 2V2 
and Camp 3, 1,980 m, Dudley 10633A (US); La Convencion, Camp 2 J /z on sharp 
ridge, ca. 1,950 m, ca. 3 A way to Camp 3, Dudley 10524 (US); Paucartambo, 
Cordillera de Tres Cruces, 3,600 m, Vargas 2231 (F); Pillahuata, Cerro de Cusil- 
luyoc, forested cliff above Rio Pillahuata, 2,300-2,400 m, Pennell 14002 (F, US). 

52. Mikania ochroleuca B. L. Robinson, Contr. Gray Herb. 80: 38. 1928. TYPE: 
Brasil, Amazonas, Rio Acre, Seringal Sao Francisco, July 1911, We 9886 (K, 
holotype, not seen, GH, photo & fragm.). 

Twining vines; stems terete, minutely crisped puberulent, internodes to 16 cm. Leaves 
deltoid-ovate, 7-8 cm long and 5-6.5 cm wide, bases hastate, 3-5-nerved from the base, 
apices acuminate, basal lobes acute to subacute, glabrous above and below, margins 
entire to undulate; petioles 3-3.5 cm long. Inflorescence paniculate, 7-13 cm high and 
6-10 cm wide, ultimate branchlets 1-2 cm long with spicately arranged heads, puberulent. 
Heads sessile to shortly pedicellate, 5-7 mm long; exterior bracts linear-lanceolate, 1-1.5 
mm long, lightly puberulent, uninervate, borne at top of pedicel; involucral bracts 
oblanceolate-oblong, ca. 4 mm long, 3-5-nerved, glabrate, glandular-atomiferous with 
somewhat orange glands, apices acute to obtuse, bases acute; corollas 3 mm long, 
whitish-yellow, tube ca. 2 mm long, throat salverform, ca. 0.5 mm long, teeth deltoid- 
ovate, ca. 0.5 mm long, recurved. Achenes ca. 2 mm long, black with white ribs, glandu- 
lar; pappus bristles 30-40, white, ca. 2.5 mm long. 

Mikania ochroleuca was previously known only from the type. It is similar in 
many respects, according to Robinson (1928), to M. hastifolia of Rio de Janeiro, 
Brazil, a plant with larger heads and achenes 2x as long as M. ochroleuca. 

CUZCO. Quispicanchis: entre Inambari y 15 Mil, 500-650 m, Vargas 16528 
(US). 

53. Mikania oreopola B. L. Robinson, Contr. Gray Herb. 64: 16. 1922. TYPE: 
Peru, St. Gavan, July 1852, Lechler2315 (K, holotype, not seen; GH, isotype). 

Slender twiners; stems somewhat hexagonal, costate, hollow, glabrous to slightly pu- 
berulent, internodes 12-17 cm long. Leaves elliptic to ovate to ca. 20 cm long and 10 cm 
wide, margins more or less entire, apices acuminate, bases narrowed to cuneate, upper 
surfaces glabrous, 5-nerved from near the base, secondary veins prominent, arching to 
the apex and somewhat parallel to the margins, reticulating veins prominent, exserted, 
lower surfaces glabrous to weakly puberulent; petioles to ca. 2 cm long, glabrate. In- 
florescence spicate; branchlets terete, costate, glabrous. Heads ca. 5 mm long; exterior 
bracts lance-linear, ca. 0.7 mm long; involucral bracts oblong, 2.5-3 mm long, glabrous, 
apices rounded; corollas semifunnelform, ca. 2.5-3 mm long, tube 1-1.2 mm long, throat 
0.7-1 mm long, teeth deltoid, ca. 0.5 mm long, glandular. Achenes ca. 1.8 mm long, 
glandular toward the summit, black, ribs white; pappus bristles ca. 40, white, ca. 2.5 mm 
long, barbellate. 

Mikania oreopola is known at present only from Peru. 

AMAZONAS. Quebrada Huampani, Rio Cenepa, 700-800 ft, Ancuash 126 
(IBE, MO). CUZCO. Quispicanchis: Entre 15 Mil & Sn. Lorenzo, 700 m, Vargas 



MACBRIDE: FLORA OF PERU 41 

U860 (US). HUANUCO. Huanuco: Cerca Tingo Maria, entre Huanuco y 
Pucallpa, 600-700 m, Ferreyra 918 (US, USM). JUNlN. La Merced, Hacienda 
Schunke, ca. 4,000 ft, Macbride 5731 (F, GH, US). LORETO. Boqueron Padre 
Abad, 470 m, Woytkowski 34419 (F, MO). SAN MARTfN. Huallaga, Muna, 
2,300-2,400 m, Weberbauer 6713 (F, GH, US). 

54. Mikania parvicapitulata Hieron., Hot. Jahrb. Syst. 40: 392. 1908. TYPE: 
Peru, Huanuco, Huamalies, mountains to S of village of Monzon, 2,500-2,900 
m, Weberbauer 3405 (B, holotype, destroyed, F, GH, photos). 

Twining shrubs; stems terete, at first rusty or dark villous, internodes 1-5 cm long. 
Leaves broadly elliptic, at most 4.5 cm long and 2.5 cm wide, margins entire, apices 
minutely toothed, mucronate, bases acute, upper and lower surfaces glandular-punctate, 
pinnately veined, with 2 pairs of lateral nerves beginning within 1 cm of base; petioles 
0.5-1 cm long. Inflorescence a panicle with heads ultimately disposed in racemes; 
pedicels 1.5-3 mm long. Heads small for genus (less than 4 mm?); exterior bracts linear, 
ca. 1.5 mm long; involucral bracts oblong, ca. 2.5 mm long, thickish, rather densely 
glandular, 1-nerved, villous on dorsal surface, apices subtruncate; corollas ca. 3 mm long, 
whitish-yellow, with globular sessile glands, tube ca. 1 mm long, throat turbinate, ca. 1.5 
mm long; corolla teeth deltoid-ovate, ca. 0.5 mm long. Achenes (immature) ca. 1 mm long, 
sessile glands on angles; pappus bristles 35-40, yellowish-white. 

Mikania parvicapitulata is known only from the type and no specimen was 
seen. The description given here is condensed from that of Robinson (1922) and 
characters discernible from photos of the type. The inflorescence of this species 
suggests the Bolivian M. dioscoreoides in head size, denseness, and disposition. 

55. Mikania parviflora (Aubl.) Karst., Deutsche Fl. 1061. 1883. 

Eupatorium parviflorum Aubl., PI. Guian. 2: 797. 1775. TYPE: not seen. 
. amarum Vahl. Symb. 3: 93. 1794. TYPE: not seen. 
Mikania amara (Vahl) Willd., Sp. PI. 3: 1744. 1803. 

M. loretensis B. L. Robinson, Contr. Gray Herb. 64: 15. 1922. TYPE: Peru, Loreto, 
Maynas, Iquitos, Ule 6338 (K, holotype, not seen, GH, photo & fragm.). 

Lianas; stems terete, finely puberulent, internodes to 20 cm or more. Leaves ovate, to 
ca. 15 cm long and 11 cm wide, margins entire, apices acute to acuminate, bases rounded 
to subcordate, texture somewhat chartaceous-coriaceous, upper surfaces glabrate to 
lightly puberulent, veins prominent, exserted, pinnate; petioles to ca. 5 cm long, terete, 
glabrous to puberulent. Inflorescence a corymb, trichotomously branching, ca. 10 cm in 
diameter; branchlets terete to irregularly angled, puberulent. Heads sessile, ternately 
disposed, 9-10 mm long; exterior bracts spatulate-elliptic, somewhat petiolate, ca. 3.5 mm 
long, apices acute to rounded, bases cuneate, borne at summit of pedicel; involucral 
bracts oblong, 6-7 mm long, glabrate to puberulent (especially at apex), apices rounded, 
bases somewhat swollen; corollas ca. 4-4.5 mm long, white to lilac in color, tube 1.5-1.8 
mm long, throat cylindric to narrowly campanulate, 2.2-2.5 mm long, teeth deltoid, ca. 0.3 
mm long, glandular. Achenes ca. 4.0 mm long, somewhat greenish to brownish; pappus 
bristles ca. 50, carneous in age, 4.5-5 mm long, barbellate. 

Mikania parviflora of northern South America is nomenclaturally the oldest 
name of a difficult group of closely related plants with sessile, ternately ar- 
ranged heads. The plant is characterized by its ovate leaves with pinnate vena- 
tion, somewhat petiolate exterior bract, and corolla teeth much shorter than the 
length of the throat. 

AMAZONAS. Quebrada Wampushik entsa, monte, 940 m, Kayap 958 (MO), 
Bagua: Valley of Rio Maranon near Cascadas de Mayasi, about km 278 of Mara- 
non Road, Wurdack 1928 (GH, US). LORETO. Balsapuerto, 220 m, Klug 3060 (F), 



42 FIELDIANA: BOTANY 

Rio Paranapura above Yurimaguas, Croat 17939 (MO), along Rio Huallaga in 
vicinity of Lagunas, Croat 17861 (MO), Gamitanacocha, Rio Mazan, 100-125 m, 
/. M. Schunke 29 (GH, MO, USM) & 204 (MO, USM), Iquitos, Ule 6235 (GH), 
Iquitos, 100 m, Killip & Smith 27355 (GH, US), Mishuyacu, near Iquitos, 100 m, 
Klug 837 (F, US), along Rio Amazon S of Iquitos, Croat 19307 (IBE), Carretera de 
San Jose across Rio Itaya, Rimachi 737 (IBE, NATC), Santa Ana on the upper Rio 
Nanay, Williams 1223 (GH), Rio Nanay, vicinity of Santa Maria de Nanay, 120 
m, Gentry & Ayala 15564 (MO), Boqueron del Padre Abad, 470 m, Woytkowski 
s.n. (MO, USM). SAN MARTfN. Tarapoto, Spruce 4118 (GH), Mariscal Caceres: 
Tocache Nuevo, Puerto Pizana, 350 m, Schunke V. 6955 (MO). 

56. Mikania pendula Holmes & McDaniel, Phytologia 37: 472. 1977. TYPE: 
Peru, Cusco, Convencion, Quellomayo to Lucomayo, 2,800 m, July 26, 1944, 
Vargas 4485 (F, holotype). 

Slightly woody lianas; stems terete, striate-costulate, glabrous to sparingly puberulent, 
intemodes to 12 cm long. Leaves ovate, to 10 cm long and 5 cm wide, margins serrate, 
teeth ca. 1 cm apart, apices attenuate, bases subcordate, upper surfaces glabrous, 5-7- 
nerved from the base, tertiary veins prominent, lower surfaces glabrate, often glandular- 
atomiferous; petioles ca. 3.5 cm long, glabrous. Inflorescence paniculate, ultimate 
branches bearing spicately arranged heads; branchlets irregularly angled to flattened, 
slightly pilose. Heads 5-6 mm long; exterior bracts elliptic, ca. Vz the length of the in- 
volucral bracts, glabrate, somewhat ciliate toward the rounded apex; involucral scales 
oblong, 3-4.5 mm long, glabrate to puberulent, margins entire, apices obtuse, bases 
slightly enlarged; corollas narrowly funnelform, 3-4 mm long, glandular, tube 1-2 mm 
long, throat gradually expanded, 1-2 mm long, teeth triangular, ca. 0.5 mm long. Achenes 
ca. 2 mm long, greenish; pappus bristles 35-40, white, ca. 3 mm long, barbellate. 

Mikania pendula is known only from Cuzco, Peru. The ovate leaves with 
subcordate bases, palmate veins, and serrate margins provide ample distinction 
from all known Peruvian, Brazilian, or Bolivian species of Mikania with spi- 
cately disposed heads. 

CUZCO. Huallpacunca: Pillahuata, Paucantambo, 2,800-3,100 m, Vargas 1917 
(GH). 

57. Mikania psilostachya DC., Prodr. 5: 190. 1836. TYPE: Peru, Poeppig 2344 
(G-DC, holotype, not seen, IDC microfiche 800. 818:11.3, F, photo). Figure 7. 

M. scabra DC, Prodr. 5: 190. 1836. TYPE: French Guiana, Cayenne, Patris (G-DC, 

holotype, not seen, IDC microfiche 800. 818: II. 5, F, photo). 
M. racemulosa Benth., Ann. Nat. Hist. 2: 109. 1839. TYPE: British Guiana, Schomburgk 

480 (K, holotype, not seen; F, US, isotypes). 

M. consanguinea Gardner, London J. Bot. 6: 448. 1847. TYPE: not seen. 
M. psilostachya DC. var. scabra (DC.) Baker in Mart., Fl. Bras. 6: 265. 1876. 
M. psilostachya DC. var. racemulosa (Benth.) Baker in Mart., Fl. Bras. 6: 265. 1876. 
M. lima Schlecht in Mart., Fl. Bras. 6: 700. 1876. TYPE: not seen. 
M. moyabambensis Hieron., Bot. Jahrb. Syst. 40: 393. 1908. TYPE: Peru, Amazonas, 

Moyabamba, Weberbauer 4476 (B, holotype, not seen, F, photo; GH, isotype). 
M. karuaiensis Badillo, Bol. Soc. Venez. Ci. Nat. 10: 296. 1946. TYPE: Venezuela, 

Bolivar, Santa Teresita de Kavanayin, Gran Sabana, Steyermark 59372 (YEN, 

holotype, not seen; F, isotype). 

Vigorous shrubby twiners; stems terete to obscurely quadrangular, solid, striate, 
densely rusty velvety, internodes to 20 cm or more. Leaves elliptic to ovate, to 13 cm long 
and 5 cm wide, pinnately nerved, reticulating veins prominent, exceedingly scabrous 
above, below usually puberulent and densely glandular-atomiferous, bases acute to 
rounded, apices acute to attenuate, margins entire, or undulate or sometimes prominently 



MACBRIDE: FLORA OF PERU 



43 



serrate; petioles 1-3 cm long, densely rusty puberulent. Inflorescence paniculate with 
elongate spicate to racemose branches, puberulent with curved pubescence. Heads 8-11 
mm long, sessile to pedicellate (up to 6 mm long); exterior bracts linear-lanceolate, at base 
of pedicel, puberulent; involucral bracts oblong with acute apices, puberulent, somewhat 
glandular, 5-6 mm long; corollas ca. 5 mm long, white, tube short, ca. 1 mm long, throat 
turbinate-subcylindric, ca. 4 mm long, teeth triangular, ca. 0.5 mm long. Achenes ca. 3 
mm long, glandular; pappus bristles ca. 40, white, 5-6 mm long, barbellate. 

Mikania psilostachya occurs from Panama throughout South America to 
Bolivia and is widespread in Peru, to ca. 1,500 m elevation. As in the case of 
many species of Mikania with wide distributions, this species exhibits great 
variation, that when studied for limited areas, may appear ample for specific 
delimitation. However, when this "complex" is examined as a whole, inter- 
mediates are encountered and it becomes apparent that there is no justification 
for retention of many of the proposed species. This species is readily identified 
by its scabrous leaves, large paniculate inflorescence with spicately or 
racemosely disposed heads, rusty pubescence of the stems, and the glandular 
nature of the involucral bracts and achenes. 

AMAZONAS. Bagua: near Campamento Stte. Montenegro, km 280-284 of 
Maranon Road, 450-600 m, Wurdack 1873 (US), al lado de Quebrada Sasa, 600 ft, 




2 nun 



4 cm 



FIG. 7. Mikania psilostachya DC. A, habit; B, head; C, flower. (After McDaniel & Rimachi 
22025 [IBE].) 



44 FIELDIANA: BOTANY 

Kayap 995 (MO), trail N of Huampani, 600-800 ft, Berlin 1844 (IBE), 
Moyobamba, 890 m, Woytkowski s. n. (MO, USM), 1,200 m, Noriega s. n. (MO, 
USM). JUNlN. Colonia Perene, 680 m, Killip & Smith 25004 (GH, US), E of 
Quimiri Bridge, near La Merced, 1,100 m, Killip & Smith 23844 (GH, US), Pichis 
Trail, San Nicolas, 1,100 m, Killip & Smith 26071 (GH, US), San Ramon, 900- 
1,300 m, Killip & Smith 24774 (GH, US). LORETO. Boqueron Padre Abad, 470 m, 
Woytkowski 34365 (F, MO), Coronel Portillo: Boqueron Padre Abad, entreTingo 
Maria y Pucallpa, 470 m, Woytkowski s. n. (US), Florida, Rio Putumayo, at 
mouth of Rio Zubineta, 180 m, Klug 2238 (F, GH, MO, US), Gamitanacocha, Rio 
Mazan, 100-125 m,/. M. Schunke 388 (F, GH, US), Rio Putumayo near Colombia 
boundary, Klug 1618 (F, GH, MO, US), Yurimaguas on the lower Rio Huallaga, 
Williams 7822 (GH), Iquitos, 120 m, Williams 3634 & 7973 (both F, GH), Iquitos, 
Versailes, 100-110 m, McDaniel 16825 (IBE, MO, NATC), Iquitos, Williams 1375 
(F, GH), Mishuyacu, near Iquitos, 100 m, Klug 331 (F, US). SAN MARTIN. 
Mishquiyacu, NE of Moyobamba, 1,800 m, Woytkowski 35338 (F, MO, UC), 
Puerto Huicte (Putante), 500-600 m, Ferreyra 4404 (MO, US, USM), San Roque, 
1,350-1,500 m, Williams 7504 (F), Saposa, 400 m, Woytkowski 7254 (GH), 
Tarapoto, We 6781 (GH), Tarapoto, 830 m, Woytkowski 35044 (F, MO, USM), 
Mariscal Caceres: Tocache Nuevo, camino a Shunte, al este del Puente de Palo 
Blanco, 500-800 m, Schunke V. 7371 (IBE, MO), Nichiza, Sanja Seca de Tipishea, 
8 km de Progresso, Schunke V. 7920 (MO), Tocache Nuevo, Fundo Melodia, 
Schunke V. 7464 (MO), Tocache, Huallaga, Poeppig 1888 (F, GH), Tocache, entre 
Uchiza y Juanjui, 400-450 m, Ferreyra 447 (MO, USM). 



58. Mikania pycnadenia B. L. Robinson, Contr. Gray Herb. 64: 18. 1922. TYPE: 
Peru, San Martin, Tarapoto, Spruce s. n. (K, holotype, not seen, GH, photo & 
fragm.). 

Branching herbaceous twiners; stems terete, tawny tomentulose, internodes to 12 cm 
long. Leaves ovate, to ca. 6 cm long and 3 cm wide, margins revolute, apices acuminate, 
bases rounded, upper surfaces puberulent, lightly covered with sessile orange globular 
glands, 5-nerved from near the base, lower surfaces pubescent, densely covered with 
sessile orange globular glands, prominently reticulate, veins exserted; petioles to 4 mm 
long, terete, tomentulose. Inflorescence a somewhat dense corymb, conspicuously brac- 
teate, ca. 7 cm wide and 4 cm long. Heads sessile, arranged in 3's, 9-10 mm long; exterior 
bracts ovate; involucral bracts oblong, ca. 6.5 mm long, puberulent, glandular, apices 
rounded; corollas ca. 4.5 mm long, tube gradually expanded into the funnelform throat, 
teeth broadly deltoid, ca. 0.5 mm long, hispidulous at the apex. Achenes ca. 4.3 mm long, 
pale brown; pappus bristles ca. 65, fulvous, 4.5-5 mm long, barbellate. 

Mikania pycnadenia is known only from the type. The above description was 
compiled from that of Robinson (1922) and items obtainable from a photo and 
fragments of the type. This species is well marked by its glandular leaves and 
petioles less than 5 mm long. 

59. Mikania rimachii Holmes & McDaniel, Phytologia 41: 195. 1979. TYPE: 
Peru, Loreto, Maynas, Dtto. Indiana, ca. 3 km up Rio Amazonas from Indiana 
on opposite bank, May 26, 1978, McDaniel, Rimachi, Holmes & Bruza 21646 
(IBE, holotype; AMAZ, F, FSU, GA, K, MO, NATC, NY, RB, RSA, TENN, US, 
USM, VDB, isotypes). Figure 8. 

M. scandens (L.) Willd. var. alata Baker in Mart., Fl. Bras. 6: 250. 1876. TYPE: Brasil, 
Santarem, Spruce 983 (F, isorype) notM. alata (G. F. Mey.) DC., Prodr. 5: 197. 1836. 



2 nun 



2 nun 




4 cm 



FIG. 8. Mikania rimachii Holmes & McDaniel. A, inflorescence; B, head; C, flower; D, 
habit. (After McDaniel et al. 21646 [IBE].) 



45 



46 FIELDIANA: BOTANY 

Lianas; stems terete, weakly striate, glabrate, internodes to 11 cm or more. Leaves 
broadly ovate, to ca. 14 cm long and 10 cm wide, margins entire, apices acuminate, bases 
shallowly cordate, upper surfaces glabrous, palmately 5-7-nerved, lower surfaces sparsely 
muricate, lighter than above, often purple; petioles to ca. 6 cm long, glabrate. In- 
florescence a somewhat cymose corymb, to ca. 6.5 cm long and 11 cm in diameter; 
branchlets angular, winged, lightly puberulent, reddish; pedicels ca. 4 mm long. Heads 
6.5-7.5 mm long; exterior bracts lanceolate to ovate, 2.5-3 mm long, glabrate to lightly 
puberulent, often reddish, apices acute to acuminate, borne at the summit of the pedicel; 
involucral bracts linear-oblong, ca. 5.5-6.5 mm long, glabrate to sparsely puberulent 
(especially at the base), with 1-3 somewhat obscure but visible nerves, stramineous in 
color, apices acute; corollas ca. 3.5 mm long, tube ca. 1.7 mm long, throat campanulate- 
funnelform, 0.7-0.8 mm long, teeth 1-1.1 mm long, about the same length to slightly 
longer than the throat, triangular. Achenes ca. 3 mm long, glabrous, greenish (slightly 
immature), ribs with somewhat narrow subherbaceous wings; pappus bristles ca. 32, 
thin, white, ca. 3.5 mm long, barbellate. 

Mikania rimachii closely resembles M. micrantha but may be distinguished by 
its larger heads of ca. 7 mm long, larger involucral bracts over 5 mm long and 
corolla teeth about the same length as the throat. Mikania micrantha has heads 6 
mm or less long, involucral bracts 4 mm or less long, and corolla teeth distinctly 
shorter than the throat. Experience in the field has shown that this species may 
be distinguished at a glance, even when sterile, by its broadly ovate leaves with 
shallowly cordate bases and often purplish lower surfaces. For a detailed dis- 
cussion of the nomenclature of this species, see Holmes & McDaniel (1979). 

LORETO. Maynas: Dtto. Iquitos, Rio Amazonas SE of Iquitos across from 
Padre Isla, near Sacarita de Yanayacu, McDaniel & Rimachi 18963 (IBE, NATC); 
Iquitos, Ushpa-Cana, across Rio Itaya, 100-110 m, McDaniel 10879 (AMAZ, IBE, 
NATC, RB, USM); Dtto. Mazan, Rio Napo near Mazan. 110 m, Mexia 6467 (F, 
GH, MICH, MO, NO, NY, PH, TEX, UC), Dtto. Las Amazonas, Rio Amazonas 
just above Orellana, Rimachi 4423 (AMAZ, F, FSU, GA, IBE, MO, NATC, NY, 
RSA, SMU, US, USM, VDB). 

60. Mikania rugosa B. L. Robinson, Contr. Gray Herb. 61: 20. 1920. TYPE: 
Peru, Puno, Ramospata, between Sandia and Chunchusmayo, 2,400-2,500 m, 
July 27, 1902, Weberbauer 1323 (B, holotype, destroyed, GH, photo & fragm.). 

Twining shrubs; stems somewhat 6-angled, densely and sometimes retrorsely tawny 
tomentose, internodes to 12 cm long. Leaves narrowly ovate, to ca. 11 cm long and 5.5 cm 
wide, margins entire to remotely and obscurely cuspidate-denticulate, apices attenuate, 
bases shallowly cordate, upper surfaces subcoriaceous, finely fulvous-pilose, bullate- 
rugose, 3-7-nerved from the base, lower surfaces densely fulvous-tomentose; petioles to 
ca. 2.5 cm long, terete, tawny- tomentose, opposite petioles connate by a transverse fold or 
ring. Inflorescence a panicle, to ca. 5 dm long and 2.5 dm in diameter; branchlets tawny- 
tomentose; pedicels 0.5-5 mm long, tawny-tomentose. Heads ca. 6 mm long; exterior 
bracts lance-oblong, to ca. 4 mm long, tomentose, borne at the summit of the pedicel; 
involucral bracts oblanceolate-oblong, to ca. 4 mm long, tawny-tomentose, apices sub- 
acute; corollas ca. 3.5 mm long, whitish, tube ca. 1.5 mm long, appressed hirtellous 
toward the summit, throat broadly campanulate, ca. 1 mm long, teeth ovate-lanceolate, 
ca. 1 mm long. Achenes ca. 3 mm long; pappus bristles ca. 30, whitish, ca. 4.5 mm long, 
barbellate. 

Mikania rugosa is known only from the type material. It is very near to M. 
banisteriae in characters of inflorescence, corolla, pubescence, etc., but may be 
distinguished by its somewhat hexagonal stems and densely tawny- 
tomentulose involucral bracts. Mikania banisteriae has terete stems and glabrate 
involucral bracts. Much of the above description is from Robinson (1922). 



MACBRIDE: FLORA OF PERU 



47 



61. Mikania shushunensis Holmes & McDaniel, Phytologia 45: 298. 1980. 
TYPE: Peru, Loreto, Maynas, Dtto. Iquitos, Carretera de Zungaro Cocha near 
Quebrada de Shushuna, September 20, 1978, Rimachi 3984 (IBE, holotype; 
AMAZ, F, NATC, US, USM, isotypes). Figure 9. 

Lianas; stems terete, costate, glabrous, pithy, internodes to ca. 7 cm long. Leaves 
lanceolate to ovate, to 10 cm long and 4.5 cm wide, apices attenuate, bases cuneate, 
margins entire, pinnately nerved, surfaces glabrous, lower surfaces prominently re- 
ticulate; petioles ca. 1.5 mm long, glabrous. Inflorescence a panicle, to ca. 15 cm long and 
7cm in diameter; branchlets puberulent. Heads somewhat racemosely disposed, ca. 6mm 
long; exterior bracts linear, puberulent, ca. 3 mm long, bome at the base of the pedicel; 
involucral bracts lanceolate, ca. 5 mm long, glabrate, apices somewhat pointed to 
rounded, puberulent; corollas white, ca. 4 mm long, tube gradually expanding into and 
scarcely distinguishable from the throat, teeth lanceolate, ca. 1.2 mm long. Achenes (im- 
mature) ca. 1.8 mm long; pappus bristles 30-35, ca. 4.5 mm long, brownish near the base, 
gradually turning whitish and slightly enlarged toward the tips, scabrid. 

Distinguishing characteristics of Mikania shushunensis include its paniculate 
inflorescence with heads somewhat racemosely disposed, linear exterior bracts 



= 2 mm 




FIG. 9. Mikania shushunensis Holmes & McDaniel. A, habit; B, head; C, flower; D, leaf. 
(After Rimachi '3984 [IBE].) 



48 FIELDIANA: BOTANY 

borne at the base of the pedicel, and bicolored pappus, the bases brownish, the 
tips white. This species is similar to M. iquitosensis. Both species appear to turn 
dark brown upon drying. The former has leaves with cuneate bases and a 
corolla with tube and throat scarcely distinguishable, while the latter has leaves 
with acute to obtuse bases and a campanulate corolla throat easily distinguish- 
able from the tube. 

LORETO. Maynas. Dtto. Iquitos. Carretera de Pena Negra, ca. 11 km from 
Quisto Cocha, on trail to Rio Itaya, sandy uplands, September 30, 1980, Rimachi 
5333 (AMAZ, F, FSU, GA, IBE, MO, NATC, NY, RB, TEX, US, USM). 

62. Mikania simpsonii Holmes & McDaniel, Phytologia 37: 473. 1977. TYPE: 
Peru, Loreto, Maynas, Dtto. Alto Nanay, trail near Santa Maria de Nanay, 
March 4, 1968, Simpson 781 (US, holorype, F, isotype). 

Lianas; stems terete, pithy, weakly puberulent, soon glabrate, internodes ca. 6.5 cm 
long. Leaves verticillate (3 per node), ovate, to ca. 14 cm long and 6.5 cm wide, margins 
entire, apices attenuate, bases acute to obtuse, upper surfaces dark green, glabrous, 
pinnately veined, with 2-3 pairs of veins prominent and arching toward the apex, lower 
surfaces glabrous, medium green, reticulate-alveolate, with at least the prominent veins 
being darkened; petioles ca. 2.5 cm long, thickish. Inflorescence a lax panicle with the 
heads ultimately disposed in open spikes; branchlets irregularly angled to terete, pu- 
berulent, with linear bracts near the base, 0.5-1.5 cm long. Heads ca. 6.5 mm long, sessile; 
exterior bracts lance-ovate, ca. Va the length of the involucre, glabrate, apices acuminate; 
involucral bracts oblong, ca. 4 mm long, glabrate, apices rounded, white inside; corollas 
ca. 3 mm long, tube ca. 1.5 mm long, throat abruptly expanded, ca. 0.5 mm long, teeth 
lanceolate, ca. 1.0 mm long. Achenes ca. 2.5 mm long, glabrous, dark brown, ribs white, 
distinctly narrowed toward the apex; pappus bristles 33-35, light cameous, ca. 3.5 mm 
long, barbellate, thickened at the tips. 

Mikania simpsonii is known from northeastern Peru and adjacent Colombia. It 
is the only known Peruvian species of Mikania with whorled leaves. 

AMAZONAS. Quebrada Huampani, Rio Cenepa, 700-800 ft, Ancuash 126 
(IBE). LORETO. Balsapuerto, 220 m, Klug 2960 (F, GH, MO, US) (paratype of M. 
klugii B. L. Robinson). SAN MARTlN. Mariscal Caceres, Tocache Nuevo, 
Fundo Gran Chaparral del Sr. Carlos Arevala, 500 m, Schunke V. 8665 (MO). 

63. Mikania speciosa DC., Prodr. 5: 196. 1836. TYPE: Peru, Haenke (G-DC, 
holorype, not seen, IDC microfiche 800. 820: 1.6, GH, photo). 

Suffruticose lianas; stems terete, glabrate to puberulent, internodes to ca. 16 cm or 
more. Leaves broadly ovate, to ca. 15 cm long and 11 cm wide, margins entire, apices 
acute, bases rounded, cuneately decurrent upon petiole at insertion, surfaces somewhat 
scabrid, pinnately nerved from near the base, prominently reticulate; petioles to ca. 3-4 
cm long, scabrid-puberulent. Inflorescence a trichotomous corymb, to ca. 10 cm or more 
in diameter, heads disposed in 3's at the tips of the branches; branchlets irregularly terete, 
puberulent. Heads 10-12 mm long, sessile; exterior bracts linear-lanceolate, ca. 3 mm 
long, puberulent; involucral bracts oblong, 7-8 mm long, puberulent, glandular, apices 
rounded to obtuse; corollas ca. 5 mm long, purplish, tube ca. 3 mm long, throat ca. 0.7mm 
long, the teeth oblong, ca. 1.3 mm long. Achenes ca. 3.5 mm long, dark grayish; pappus 
bristles ca. 65, carneous (in age), ca. 5 mm long, barbellate. 

Mikania speciosa occurs to about 2,400 m elevation in Bolivia and Peru. 

AMAZONAS. Chachapoyas, Quebrada Molina 5 km below Chachapoyas, 
2,200-2,400 m, Wurdack 7770 (F, US). CUZCO. San Miquel, Urubamba Valley, 
1,800 m, Cook & Gilbert 947 (US); Villcabamba, hacienda on Rio Chinchao, 6,000 



MACBRIDE: FLORA OF PERU 49 

ft, Macbride 5190 (GH, US). SAN MARTlN. Rio Huallaga Canon, below Santo 
Domingo, ca. 4,000 ft, Macbride 4198 (F, GH, US). 

64. Mikania stereodes B. L. Robinson, Contr. Gray Herb. 77: 60. 1926. TYPE: 
Peru, Cuzco, hacienda on Rio Chinchao, July 17-26, Macbride 5163 (F, 
holotype). 

Woody twiners; stems terete, multicostate, fulvous, somewhat scabrid, internodes to 20 
cm long. Leaves ovate, ca. 10 cm long and 6 cm wide, margins entire, apices acuminate, 
bases obtuse, upper surfaces scabrid, pinnately nerved, the major nerves strigose, lower 
surfaces puberulent, the major nerves strigose, prominently reticulate, minutely dark 
punctate; petioles to 1.5 mm long, strigose. Inflorescence thyrsoid, ca. 25 cm long and 10 
cm wide; branchlets terete, strigose; pedicels to ca. 3 mm long, terete, strigose. Heads 8-9 
mm long; exterior bracts lanceolate-linear, ca. 2 mm long, pilose, especially toward the 
margins; involucral bracts oblong, ca. 4 mm long, outer strigulose, inner glabrate, some- 
what dark punctate, apices obtuse to rounded; corollas ca. 5 mm long, green, remotely 
pilose, tube ca. 1 mm long, throat subcylindric, ca. 3 mm long, teeth triangular, ca. 1 mm 
long. Achenes ca. 4 mm long, light olivaceous; pappus bristles ca. 30, white, ca. 4 mm 
long, barbellate. 

Mikania stereodes, known only from the type, resembles M. psilostachya in 
having pinnately veined leaves, scabrid leaf surfaces and subcylindric corollas 
with a very short tube. Mikania psilostachya is easily distinguished by its 
spikelike to racemose inflorescence, rather than the panicle of M. stereodes. 

65. Mikania stuebelii Hieron., Bot. Jahrb. Syst. 21: 333. 1895. TYPE: Colombia, 
near El Tambo on Mt. Munchique not far from Popayan, 3,000 m, Stuebel 320 
(B, holotype, destroyed). 

M. stuebelii var. latifolia Hieron., Bot. Jahrb. Syst. 28: 577. 1901. TYPE: Colombia, 

Cauca, western Popayan Andes, 2,500-2,900 m, Lehmann 5970 (K, holotype, not seen; 

F, isotype). 
M. meridensis Badillo, Bol. Soc. Venez. Ci. Nat. 9: 134. 1944. TYPE: Venezuela, Merida, 

entre Santa Cruz de Mora y Canagua, Badillo 609a (VEN, holotype, not seen). 
M. larensis Badillo, Bol. Soc. Venez. Ci. Nat. 10: 298. 1946. TYPE: Venezuela, Lara, entre 

Santo Domingo y Los Quebraditos, 2,430-2,475 m, February 8, 1944, Steyermark 55380 

(VEN, holotype, not seen; F, isotype). 
M. stuebelii var. larensis (Badillo) Steyermark, Fieldiana, Bot. 28: 661. 1953. 

Climbing vines; stems terete, striate-sulcate, glabrate to puberulent, internodes to 4 cm 
long. Leaves lance-ovate, to ca. 7 cm long and 4 cm wide, margins remotely denticulate, 
apices acuminate, bases cuneate to obtuse, surfaces glabrous, pinnately 4-5-nerved; 
petioles ca. 1 cm long, glabrate. Inflorescence thyrsoid, heads arranged in 3's at the tips of 
the branches or more often somewhat spicately in 5's; branchlets puberulent. Heads ca. 9 
mm long, sessile; exterior bracts linear, 2-3 mm long; involucral bracts linear-oblong, 5-6 
mm long, glabrous, ciliolate, apices somewhat rounded; corollas 4-4.5 mm long, tube ca. 
1.5 mm long, throat subcylindric-campanulate, 2.5-3 mm long, teeth triangular, ca. 0.5 
mm long. Achenes ca. 4 mm long, dark, puberulent at summit; pappus bristles 40-50, 
white, 4-5 mm long, barbellate. 

Mikania stuebelii is found in northern South America, and is known in Peru 
from Amazonas, Cuzco, and Huanuco from 1,500-3,000 m elevation. The in- 
florescence of this species is distinctive, a panicle with heads arranged in 3's at 
the tips of the branches or more often somewhat spicately disposed with five 
heads, three of these being terminal and sessile. 

AMAZONAS. Chachapoyas, Cerros Calla Calla, E side, 7.5 km above 
Leimebamba on road to Balsas, 2,535 m, Hutchison & Bennett 4635 (F). CUZCO. 
Villcabamba, hacienda on Rio Chinchao, 6,000 ft, Macbride 5163 (US). 



50 FIELDIANA: BOTANY 

HUANUCO. SW slope of Rio Llullapichis watershed on the ascent of Cerros del 
Sira, 1,565-1,950 m, Dudley 13322 (US); Sariapampa, Woytkowski 34300 (F, MO). 

66. Mikania stygia B. L. Robinson, Contr. Gray Herb. 77: 36. 1926. TYPE: 
Bolivia, La Paz, Mapiri, 850 m, Buchtien 1847 (GH, holotype). 

Semiwoody climbers; stems terete, glabrate to puberulent, internodes to 17 cm long. 
Leaves lanceolate, to ca. 12 cm long and 2.5 cm wide, margins entire, apices caudate- 
acuminate, bases acute-obtuse, surfaces glabrous, pinnately nerved; petioles ca. 1.5 cm 
long. Inflorescence a corymb, ca. 4 cm long, 4 cm wide, heads in 3's at the tips of the 
branches; branchlets puberulent. Heads ca. 10 mm long, sessile; exterior bracts elliptic, 
ca. 5 mm long, puberulent; involucral bracts oblong, ca. 8 mm long, puberulent, nerves 
prominent, apices rounded; corollas ca. 6 mm long, glabrate, tube gradually expanding 
into and indistinguishable from the throat, teeth broadly deltoid, ca. 0.5 mm long. 
Achenes (immature) ca. 3 mm long; pappus bristles ca. 45, ca. 5 mm long, barbellate. Style 
appendages densely hirsute. 

The species is known from Peru and Bolivia. Mikania stygia and M. lanceolata 
are two species with corymbose inflorescence and sessile ternately disposed 
heads with lanceolate leaves. Mikania stygia has heads ca. 10 mm long and 
oblong involucral bracts ca. 8 mm long, while M. lanceolata has heads ca. 6 mm 
long and spatulate involucral scales ca. 3 mm long. 

LORETO. Alto Amazonas, Balsapuerto, lower Rio Huallaga Basin, 150-350 m, 
Killip & Smith 28616 (US). 

67. Mikania szyszylowiczii Hieron., Bot. Jahrb. Syst. 36: 475. 1906. TYPE: 
Peru, Cajamarca, near Tambillo, Jelski 681 (B, holotype, destroyed, F, photo, 
GH, fragm.). 

Somewhat woody lianas; stems glabrous, terete, internodes to ca. 14 cm long. Leaves 
ovate to somewhat obovate, 4-6 cm long, about 2 as wide, glabrous, pinnately nerved, 
bases cuneate, apices rounded, margins entire; petioles 1-2 cm long. Inflorescence 
paniculate; ultimate branchlets spicate. Heads 5-7 mm long; exterior bracts elliptic, ca. Vz 
the length of the involucre; involucral bracts oblong, irregularly ciliolate toward the 
rounded apex, bases slightly gibbous, 3-3.5 mm long; corollas 3-4 mm long, white, tube 
1.5-2 mm, throat campanulate-turbinate, 1-1.5 mm long, teeth triangular, ca. 0.5 mm 
long. Achenes dark brown, ca. 2 mm long; pappus bristles ca. 30, pale yellow, thickened 
toward the tips, 3-4 mm long, barbellate. 

Mikania szyszylowiczii is known from Peru and Ecuador (Espinoza 663, F). At 
present it is the only Peruvian species of Mikania with spicately disposed heads 
and with leaves with rounded apices and cuneate bases. 

AMAZONAS. Bongara, Jalca zone, 3 km S of Pomacocha, Wurdack 976 (US). 
CAJAMARCA. Jaen, Weberbauer 6114 (F, GH). HUANUCO: Carpish, above 
Acomayo, Hutchinson, Wright & Straw 5949 (F, MO, US). JUNfN. La Merced, 
Soukup 3370 (F). SAN MARTfN. Zepalacio near Moyobamba, King 3412 (F, MO). 

68. Mikania tambillensis Hieron., Bot. Jahrb. Syst. 36: 470. 1909. TYPE: Peru, 
Cajamarca, near Tambillo, Jelski 762 (B, holotype, destroyed; GH, not seen, 
US, isotypes). 

Twining or somewhat shrubby lianas; stems glabrous, subterete, internodes to ca. 7 cm 
long. Leaves ovate-oblong, to ca. 8 cm long and 4 cm wide, margins entire, apices acumi- 
nate, bases pointed to rounded, pinnately 5-nerved; petioles to ca. 2 cm long, smooth. 
Inflorescence a dense corymb, somewhat pyramidal, heads arranged in groups of 3's at 
the tips of the branches, these groups arranged in groups of 3-5, imparting a somewhat 
congested, glomerate look; branchlets angular, puberulent. Heads ca. 8 mm long, sessile; 



MACBRIDE: FLORA OF PERU 



51 



exterior bracts linear, ca. 2 mm long, ciliolate at summit; involucral bracts lance-linear, ca. 
5 mm long, apices obtuse; corollas 6-6.5 mm long, tube gradually expanded into a 
cylindric-subulate throat, teeth deltoid, ca. 0.5 mm long. Achenes immature; pappus 
bristles ca. 33, buff, ca. 6 mm long, barbellate. 

Mikania tambillensis is known only from the type. Due to the immature nature 
of the heads, much of the above description is from Robinson (1922). A very 
closely related plant is M. weberbaueri, which differs only in having more at- 
tenuate leaves. 

69. Mikania ternata (Veil.) B. L. Robinson, Contr. Gray Herb. 39: 198. 1911. 
Figure 10. 

Cacalia ternata Veil., Fl. Flum. 336. 1825. TYPE: not seen. 
Mikania dentata Spreng., Syst. 3: 422. 1826. TYPE: not seen. 

M. apiifolia DC, Prodr. 5: 202. 1836. TYPE: Brasil, Sao Paulo (P, holotype, not seen; 
G-DC, isotype, not seen, IDC microfiche 800. 823: 1-4). 




2 mm 



= 2 nun 



FIG. 10. Mikania ternata (Veil.) B. L. Robinson. A, habit; B, head; C, flower. (After 
Coleman s. n. [IBE].) 



52 FIELDIANA: BOTANY 

Twining vines; stems terete, glabrate to crisped puberulent, intemodes 4-8 cm long. 
Leaves to ca. 8 cm long and 4 cm wide, palmately divided into 3-7 segments, the segments 
lanceolate to narrowly elliptic-ovate, the margins entire to somewhat coarsely serrate- 
crenate to sometimes 3-lobed, often apiculate to end of major veins, apices acute, surfaces 
glabrate to lightly puberulent, segments pinnately veined; petioles 2-6 cm long, glabrous 
to puberulent, connected by a prominent incised stipule-like appendage. Inflorescence a 
compound corymb, ca. 15 cm long and 6 cm wide; branchlets somewhat angular, crisped 
puberulent; pedicels 0.5-1.5 mm long, terete, puberulent. Heads ca. 10 mm long; exterior 
bracts lanceolate to narrowly ovate, 5-6 mm long, glabrate, apices acute to attenuate, 
borne slightly below head; involucral bracts elliptic to narrowly ovate, ca. 8 mm long, 
glabrate, apices acute to attenuate; corollas semicampanulate, ca. 7 mm long, white, tube 
ca. 2 mm long, throat ca. 4.25 mm long, teeth obtuse, ca. 0.75 mm long. Achenes 4-5 mm 
long, brownish, ribs white; pappus bristles 80-100, white, 5-6 mm long, barbellate. 

Side-by-side comparison of the Peruvian material of M. ternata with that of 
elsewhere in its range (Brazil, Paraguay, and Argentina) revealed no differences 
worthy of taxonomic recognition. This species is apparently present as a dis- 
junct in Peru. It is readily distinguished from M. filici folia, the other Peruvian 
species of Mikania with parted leaves, by its palmately divided leaves, rather 
than pinnate, fernlike divisions. 

HUANUCO. Tingo Maria, Asplund 12107 (US); Hacienda El Aquila, cerca 
Tingo Maria, entre Huanuco y Pucallpa, Ferreyra 4180 (MO, US, USM); Tingo 
Maria, carretera Huanuco-Pucallpa, 700 m, Ferreyra 912 (MO, US, USM). 
LORETO. Divisoria, entre Tingo Maria y Pucallpa, 1,500-1,600 m, Ferreyra 1011 
(MO, US, USM). 

70. Mikania trachodes B. L. Robinson, Contr. Gray Herb. 61: 22. 1920. TYPE: 
Peru, Huanuco, Huamalies, mountains SW of Monzon, 2,500-2,900 m, July 
13, 1903, Weberbauer 3395 (B, holotype, destroyed, GH, photo & fragm.). 

Twining shrubs; stems terete, striate-costulate, strigillose-papillose, scabrous, inter- 
nodes to 12 cm or more. Leaves broadly ovate, subcoriaceous, to ca. 15 cm long and 8 cm 
wide, margins somewhat remotely callous-denticulate, apices acuminate, bases rounded, 
upper surfaces appressed tawny-puberulent, pinnately veined, lower surfaces spreading 
villous; petioles ca. 1.5 cm long, yellow tomentose, opposite petioles connected by a 
transverse ring or fold. Inflorescence a dense panicle, to 40 cm or more wide; branchlets 
villous. Heads 7-8 mm long; exterior bracts lanceolate, ca. Vz the length of the involucre, 
loosely villous, apices acute, borne at the summit of the pedicel; involucral bracts lanceo- 
late to oblong, ca. 4 mm long, villous, apices acute; corollas ca. 3.5 mm long, tube ca. 1.5 
mm long, pubescent at the summit, throat campanulate, ca. 1 mm long, teeth lance-ovate, 
ca. 1 mm long. Achenes ca. 2 mm long, olivaceous; pappus bristles ca. 40, white, ca. 3.5 
mm long, barbellate. 

Mikania trachodes is known only from Huanuco above 2,500 m. It is another 
species difficult to distinguish from M. banisteriae in characters of corollas, 
inflorescence, pubescence, etc. It was separated in Robinson's (1922) key to the 
species of Mikania of Peru from M. lanuginosa and M. ruiziana (both = M. 
banisteriae) by its very prominent interpetiolar ring or fold connecting opposite 
petioles, a condition not characteristic of M. banisteriae. In addition M. trachodes 
has a scabrous stem. With only a few fragments and photos available for direct 
comparison and study, it appears appropriate to retain M. trachodes as a distinct 
species. 

HUANUCO. Huanuco: Carpish, above Acomayo, 2,800 m, Hutchison, Wright 
& Straw 5962 (F, MO). 



MACBRIDE: FLORA OF PERU 53 

71. Mikania turbaricola Holmes & McDaniel, Phytologia 37: 474. 1977. TYPE: 
Peru, Amazonas, Chachapoyas, Jalca zone 3-6 km W of Molinopampas, 
2,200-2,450 m, July 19, 1962, Wurdack 1399 (US). 

Subshrubs to short vines; stems terete to angled, sulcate after drying, internodes to 5 
cm long. Leaves ovate, semicoriaceous, to ca. 7 cm long and 4 cm wide, margins entire to 
irregularly and coarsely dentate, the teeth remote, apices attenuate, bases truncate to 
rounded, upper surfaces glabrous, sparsely glandular-punctate; petioles ca. 1 cm long, 
grooved above. Inflorescence paniculate, to ca. 15 cm long and 6 cm wide; branchlets 
somewhat angled, crisped puberulent to pilose. Heads ca. 9 mm long; exterior bracts 
lance-linear to oblanceolate, ca. 3.5 mm long, borne slightly beneath the head, puberu- 
lent; involucral bracts lanceolate to lance-elliptic, ca. 6 mm long, puberulent, ciliolate at 
apex, apices acute; corollas ca. 4.5 mm long, white, glandular, tube ca. 1.7 mm long, 
throat semicampanulate, 2.2-2.3 mm long, teeth deltoid, ca. 0.5 mm long. Achenes (im- 
mature) ca. 3 mm long, green, glandular; pappus bristles 28-30, white, ca. 4.5 mm long, 
barbellate, thickened at the tips. 

Mikania tubaricola is a subshrub or short vine known only from the holotype 
collected above 2,000 m in a Sphagnum seepage bog. 

72. Mikania vargasii Holmes & McDaniel, Phytologia 35: 330. 1977. TYPE: 
Peru, Cuzco, Quispicanchis, 3,250 m, June 10, 1963, Vargas 14565 (US, 
holotype). 

Twining lianas; stems costate to sulcate (after drying), glabrous, internodes 6-10 cm 
long. Leaves ovate to ovate-oblong, to ca. 6 cm long and 4 cm wide, margins entire, apices 
attenuate, bases cordate, upper surfaces sparsely puberulent, dark green, 5-nerved from 
the base, lower surfaces glabrous, dark punctate with globular sessile glands, lighter than 
above; petioles to ca. 2 cm long, thin, glabrous. Inflorescence paniculate, the ultimate 
branchlets spicately disposed, ca. 20 cm long and 10 cm wide; branchlets angular, pilose. 
Heads 10-12 mm long, sessile; exterior bracts elliptic, 3-3.5 mm long, glabrate, apices 
acute; involucral bracts oblong to obovate, ca. 6 mm long, appressed-puberulent, entire, 
apices rounded to obtuse, bases slightly gibbous; corollas semisalverform, lightly glan- 
dular, ca. 6 mm long, tube ca. 4 mm long, throat ca. 1 mm long, teeth triangular, ca. 1 mm 
long. Achenes 4-4.5 mm long, dark brown to black; pappus bristles 80-85, white, 5-6 mm 
long, barbellate, gradually narrowed from base to tip. 

Mikania vargasii, known only from Cuzco above 2,000 m, is easily recognized 
by the large paniculate inflorescence with the spicately disposed heads 10-12 
mm long. 

CUZCO. Calca, Manfio, K84, 2,000-2,200 m, Vargas 15634 (US). 

73. Mikania violaefolia Cuatrecasas, Ann. Missouri Bot. Card. 52: 307. 1965. 
TYPE: Peru, Amazonas, Chachapoyas, km 416-419 of Leimebamba-Balsa 
Road, 2,900-3,100 m, July 9, 1962, Wurdack 1257 (US, holotype; F, isotype). 

Twining vines; stems angular to somewhat hexagonal, purplish, glabrous, internodes 
to 15 cm long. Leaves triangular to deltoid, to ca. 3 cm long and 3 cm wide, margins 
coarsely crenate-dentate, apices acute, bases cordate, upper surfaces somewhat rugose, 
palmately 5-7-nerved, lower surfaces glabrate-puberulent, prominently reticulate; 
petioles 1.5 cm long, glabrous, connected by a laciniate stipule-like ring. Inflorescence a 
corymb, ca. 4 cm in diameter; branchlets angular, glabrous; pedicels to 8 mm long, 
glabrate. Heads ca. 10 mm long; exterior bracts lanceolate to oblanceolate, purple tinged, 
ca. 5 mm long, glabrate to puberulent, apices acute, borne well beneath the involucre; 
involucral bracts lance-oblong, 8-9 mm long, glabrous, purplish, apices acute, base nar- 
rowly bulbous; corollas ca. 5.5 mm long, white, tube and throat indistinguishable, teeth 
triangular, ca. 1.5 mm long. Achenes ca. 3 mm long; pappus bristles ca. 100, white, ca. 5 
mm long, barbellate. 



54 FIELDIANA: BOTANY 

Mikania violaefolia, known only from Amazonas above 2,900 m, is charac- 
terized by its small (usually less than 1.5 cm long) leaves, with coarsely 
crenate-dentate margins. 

AMAZONAS. Chachapoyas: Cerros Calla Calla, 18km above Leimebamba on 
road to Balsas, 3,100 m, Hutchison & Wright 5679 (F, MO, US). 

74. Mikania vitifolia DC., Prodr. 5: 202. 1836. TYPE: Brasil, Sao Paulo (P, 
holotype, not seen; G-DC, isotype not seen, IDC microfiche 800. 823: 1.2). 
Figure 11. 

M. gracilis Schultz-Bip. ex Miq., Stirp. Surin. Sel. 187. 1850. TYPE: Surinam, Hostmann 

& Kappler 1017 (G, holotype, not seen; F, isotype). 
M. punctata Klatt, Bull. Soc. Bot. Belgique 31: 195. 1893. TYPE: Costa Rica, Pittier3434 & 

4934, syntypes, BR). 
M. boliviensis Lingelsh., Fedde Rep. 7: 251. 1909. TYPE: Bolivia, San Carlos near Mapiri, 

September 1907, Buchtien 71 (F, isotype). 
M. cryptodonta B. L. Robinson, Contr. Gray Herb. 73: 22. 1924. TYPE: Peru, Pozuzo, 

June 20-22, 1923, Macbride 4683 (F, holotype). 
M. lechleri Schultz-Bip. ex B. L. Robinson, Contr. Gray Herb. 77: 35. 1926. TYPE: Peru, 

Puno, Carabay, St. Govan, August 1854, Lechler 2480 (K, holotype, not seen, GH, 

photo & fragm.), including M. lechleri Schultz-Bip., Bonplandia 4: 54. 1856, nomen 

nudum. 
M. caustolepis B. L. Robinson, Contr. Gray Herb. 77: 49. 1926. TYPE: Peru, Cuzco, Rio 

Yanamayo, below Pillahuata, 2,000-2,300 m, May 4-5, 1925, Pennell 14070 (GH, 

holotype; F, isotype). 

M. vitifolia f. boliviensis (Lingelsh.) B. L. Robinson. Contr. Gray Herb. 104: 54. 1934. 
M. vitifolia f. cryptodonta (B. L. Robinson) B. L. Robinson, Contr. Gray Herb. 104: 55. 

1934. 
M. anzoatiguensis Badillo, Bol. Soc. Venez. Ci. Nat. 10: 301. 1946. TYPE: Venezuela, 

Anzoatequi, NE of Bergantin, March 1-2, 1945, Steyermark 61376 (YEN, holotype, not 

seen; F, isotype). 

Herbaceous to subwoody lianas; stems terete, striate to costate, fistulose in age, pu- 
berulent to pilose at nodes, internodes to 25 cm long. Leaves ovate, often with 2 divergent 
basal lobes in the lower Va of the leaf, to 20 cm long and 17 cm wide, margins entire, 
apices acuminate-attenuate, base obtuse to truncate to subcordate or cordate, surfaces 
glabrate to puberulent, usually dark glandular-punctate, palmately or subpinnately 5-7- 
nerved from or near the base; petioles to 11 cm long, glabrate to puberulent. Inflorescence 
elongated, bracteate, lax to dense panicle to 5 dm or more long; branchlets terete to 
angular, puberulent; pedicels 1-2 mm long. Heads 7-9 mm long; exterior bracts lanceo- 
late, ca. 2 mm long, puberulent, glandular; involucral bracts oblong, 4.5-5 mm long, 
glandular, margins thin, often whitish in color, nerves prominent, apices puberulent, 
obtuse; corollas ca. 5 mm long, white, glandular (especially on teeth), tube ca. 3 mm long, 
thin, throat ca. 1 mm long, teeth triangular, ca. 1 mm long. Achenes ca. 3.5 mm long, 
glandular, greenish-brown; pappus bristles 35-40, thin, ca. 5 mm long, barbellate. 

Mikania vitifolia is a widespread (southern Mexico to Bolivia and Brazil) and 
somewhat polymorphic species with leaf shape varying greatly. Mature cauline 
leaves generally have two divergent, pointed basal lobes ca. two-thirds the 
distance from the apex. Often this character is not evident, because many 
specimens have only bracteal leaves and the inflorescence. 

ANCASH. Pampayacu, hacienda at mouth of Rio Chinchao, 3,500 ft, Macbride 
5062 (F, GH, US). CUZCO. Convencion: Hacienda Portusco, 1,300 m, Vargas 
11677 (F, US), Paucartambo, Hacienda Partria, Marin 1717 (F, US). HUANUCO. 
Pozuzo, ca. 2,000 ft, Macbride 4552 (F, GH, US), Rio Huallaga Canon, below 
Santo Domingo, Macbride 4219 (F, GH, US). JUNfN. La Merced, Hacienda 



MACBRIDE: FLORA OF PERU 



55 



= 2 mm 




4 cm 

FIG. 11. Mikania vitifolia DC. A, habit; B, head; C, flower; D, leaf. (After McDaniel & 
Rimachi 22029 [IBE].) 



Schunke, ca. 4,000 ft, Macbride 5461 (F, GH, US), E of Quimiri Bridge, near La 
Merced, 800-1,300 m, Killip & Smith 23835 (F, GH, US), Rio Paucartambo Valley 
near Perene Bridge, 700 m, Killip & Smith 25313 (F, GH, US), Sanibeni, 900 m, 
Woytkowski 5038 (GH). SAN MARTiN. Valley of San Martin along Rio Shilcayo, 
1-4 mi NE of Tarapoto, ca. 1,000 ft, Belshaw 3282 (GH, MO, US), Tarapoto, 
Carretera Tarapoto-Yurimaguas, km 12-15, 2,250 ft, McDaniel & Rimachi 22019 
(AMAZ, F, IBE, MO, NY, US, USM). 



56 FIELDIANA: BOTANY 

75. Mikania weberbaueri Hieron., Bot. Jahrb. Syst. 11: 389. 1908. TYPE: Peru, 
Huamalies, Monzon, 100-1,000 m, July 23, 1903, Weberbauer 3426 (B, 
holotype, destroyed, F, GH, photos). 

M. rivularis B. L. Robinson, Contr. Gray Herb. 73: 30. 1924. TYPE: Peru, Junfn, La 
Merced, 2,000 ft, August 10-24, Macbride 5414 (F, holotype). 

Twining shrubs; stems terete, striate-sulcate, glabrous (?), intemodes to 19 cm long. 
Leaves ovate-oblong to broadly lanceolate, to ca. 13 cm long and 5 cm wide, margins 
entire, apices long attenuate, bases obtuse to somewhat cuneate, surfaces smooth, sub- 
pinnately 3-5-nerved from near the base; petioles to ca. 2 cm long. Inflorescence a panicle 
composed of rather dense glomerate corymbs, the corymbs 2-3 cm in diameter. Heads 7-8 
mm long, sessile to nearly so; exterior bract lanceolate, ca. 3 mm long; involucral bracts ca. 
5 mm long, puberulent, 5-7-nerved, apices acute to obtuse, ciliolate; corollas ca. 6 mm 
long tube gradually expanded into the throat, teeth oblong, ca. 0.5 mm long. Achenes ca. 
2 mm long; pappus bristles ca. 35-40, white, thickened at tips, obscurely barbellate. 

Mikania weberbaueri is known only from Peru. The actual type specimen was 
not seen. The description is compiled from that of Robinson (1922) and photos 
and sketches of the type. This species is uncomfortably close to M. tambillensis, 
but is distinguished from that species, which has acute to abruptly acuminate 
leaf apices, by its long attenuate leaf apices. 

From available evidence Mikania weberbaueri and M. rivularis cannot be sepa- 
rated and are to be considered conspecific. 

76. Mikania woytkowskii Holmes & McDaniel, Phytologia 37: 475. 1977. TYPE: 
Peru, Loreto, Boqueron Padre Abad, August 20, 1946, Woytkowski 34393 (F, 
holotype; MO, isotype). 

Herbaceous lianas; stems terete, costate, hollow, glabrous, internodes to 10 cm long. 
Leaves elliptic to narrowly ovate, to 7 cm long and 2.5 cm wide, margins entire, apices 
acute to acuminate, bases cuneate, upper surfaces glabrous, 3-5-nerved from the base, 
veins exserted from lower surfaces, glabrous, veins exserted; petioles to ca. 1 cm long. 
Inflorescence paniculate, the ultimate branchlets spicate; branchlets costate, puberulent. 
Heads 6-7 mm long; exterior bracts elliptic, ca. Va-Vz the length of the involucre, spar- 
ingly puberulent, apices acute, margins slightly ciliolate; involucral bracts elliptic-oblong, 
ca. 4.5 mm long, glabrate, 3-5{7)-nerved, apices obtuse; corollas funnelform, 3.5-4 mm 
long, white, tube 1.5-2 mm long, throat ca. 1.5 mm long, teeth deltoid, ca. 0.5 mm long. 
Achenes ca. 2 mm long, glandular, dark brown (immature); pappus bristles ca. 35, white, 
ca. 4 mm long, barbellate, thickened at the tips. 

Mikania woytkowskii, known only from the type, is a slender herbaceous, 
twining vine. A similar species is M. oreopola, which is distinguished by its 
ovate leaves with obtuse bases, prominent, reticulating veins, and elliptic in- 
volucral scales to 3 mm long. 




1 TUMBES 

2 PIURA 

3 CAJAMARCA 

4 AMA2ONAS 

5 LORETO 

6 SAN MARTIN 

7 LAMBAYEOUE 

8 LA LIBERTAD 

9 ANCASH 

10 HUANUCO 

1 1 LIMA 

12 PASCO 

13 JUNIN 

14 HUANCAVELICA 

15 ICA 

16 AYACUCHO 

17 APURIMAC 

18 CUZCO 

19 MADRE DE DIGS 

20 AREQUIPA 

21 PUNO 

22 MOOUEGUA 

23 TACNA 



100 200 300 



PERU 
Provinces 



UNIVERSITY OF ILLINOIS-URBANA 





30112028176045 




Field Museum of Natural History 
Roosevelt Road at Lake Shore Drive 
Chicago, Illinois 60605-2496 

Telephone: (312) 922-9410 




UNIVERSITY OF ILLINOIS-URBANA